Chapter IV: Part 4
This fragment, which was found in the bed of the same river, (see p. 16,) in Banda Oriental, with the cranium of the Toxodon, includes the parietes of the left side of the cerebral cavity, the corresponding nervous and vascular foramina, the left occipital condyle, a portion of the left zygomatic process, and, fortunately also, the left articular surface for the lower jaw. The importance of this surface in the determination of the affinities of a fossil animal has been duly appreciated, since the relations of the motions of the lower jaw to the kind of life of each animal were pointed out by Cuvier; but yet we should be deceived were we to establish, in conformity with the generalization enunciated by Cuvier,[29] our conclusion, from this surface, of the nature of the food of the extinct species under consideration; for the glenoid cavity is so shaped as to allow the lower jaw free motion in a horizontal plane, from right to left, and forwards or backwards, like the movements of a mill-stone; and, nevertheless, I venture to affirm it to be most probable, that the food of _Glossotherium_ was derived from the animal and not from the vegetable kingdom; and to predict, that when the bones of the extremities shall be discovered, they will prove the Glossothere to be not an ungulate but an unguiculate quadruped, with a fore-foot endowed with the movements of pronation and supination, and armed with claws, adapted to make a breach in the strong walls of the habitations of those insect-societies, upon which there is good evidence in other parts of the present cranial fragment, that the animal, though as large as an ox, was adapted to prey.
We perceive, in the first place, looking upon the base of this portion of skull, a remarkable cavity, situated immediately behind the tympanic bone, of nearly a regular hemispherical form, an inch in diameter (fig. 2, _b_, Pl. XVI). The superficies of this cavity appears not to have been covered with articular cartilage, for it is irregularly pitted with many deep impressions; and I conclude, therefore, that it served to afford a ligamentous attachment to the styloid element of a large _os hyoides_. With this indication of the size of the skeleton of the tongue, is combined a more certain proof of the extent of its soft, and especially its muscular parts, in the magnitude of the foramen, for the passage of the lingual or motor nerve (_c._ fig. 2 and 3). This foramen, (the anterior condyloid,) in the present specimen, is the largest of those which perforate the walls of the cranium, with the exception of the foramen magnum; it is fully twice the size of that which gives passage to the second division of the fifth nerve; its area is oval, and eight lines in the long diameter, so that it readily admits the passage of the little finger.
It is only in the Ant-eaters and Pangolins that we find an approximation to these proportions of the foramen for the passage of the muscular nerve of the tongue; and the existing Myrmecophagous species even fall short of the larger fossil in this respect. Some idea of the size of the lingual nerve, and of the organ it was destined to put in motion, may be formed, when it is stated that the foramen giving passage to the corresponding nerve in the Giraffe,—the largest of the Ruminants, and having the longest and most muscular tongue in that order,—is scarcely more than one-fourth the size.
With these indications of the extraordinary development of the tongue, we are naturally led, in order to carry out a closer and more detailed comparison of the fossil in question, to that group of mammalia in which the tongue plays the chief part in the acquisition of the food. The size, form, and position of the occipital condyle,—the magnitude of the occipital foramen, (which must here have somewhat exceeded three inches in the transverse diameter,)—the slope of the occipital surface of the cranium from below, upwards and forwards, at an angle of 60° with the base of the cranial cavity-each and all attest the close affinities of the present animal to the Edentata. More decisive evidence of the same relationship will be adduced from the organization of other parts of the cranium. The glenoid articular surface (_a_, fig. 2, Pl. XVI.) is an almost flattened plane, wider in the transverse than in the longitudinal direction; and, as in the genera _Myrmecophaga_ and _Manis_, it is not defended behind by any descending process. In its general form it resembles the glenoid cavity of _Orycteropus_ more than that of the preceding Edentates; but, in _Orycteropus_, the articulation is defended posteriorly by a descending process of the zygoma, and it is also situated relatively closer to the os tympanicum.
Had the _Glossotherium_ teeth? The extent of the temporal muscle, which is indicated by the rugged surface of the temporal fossa, and by the well-marked boundary, formed by a slightly elevated bony ridge, which extends to near the line of the sagittal suture, together with the size of the zygomatic portion of the temporal bone, and the remains of the oblique suture by which it was articulated to the malar bone, enables me to answer this question confidently in the affirmative. They will probably be found to be molar teeth of a simple structure, as in the Orycteropus.
The evidence just alluded to of the existence of an os malæ is interesting, because this bone is wanting in the Pangolins; and its rudimental representative in the true Ant-eaters does not reach the zygomatic process of the temporal bone, which consequently has no articular or sutural surface at its anterior extremity. In the presence, therefore, of the surface for the junction of the os malæ, and the consequent evidence of the completion of the zygomatic arch, we learn that the Glossothere was more nearly allied to the Armadillos and Orycterope. That its affinity to the latter genus was closer than to the Armadillos we have most interesting evidence in the form and loose condition of the tympanic bone: it is represented of the natural size at fig. 4, Pl. XVI. Through the care and attention devoted to his specimens by their gifted discoverer, this bone was preserved _in situ_, as represented at _d_, fig. 1; but it had no osseous connection with the petrous or other elements of the temporal bone, and could be displaced and replaced with the same ease as in the _Orycterope_. This bony frame of the membrana tympani, in the Glossothere, describes rather more than a semicircle, having the horns directed upwards; it has a groove, one line in breadth, along its concave margin, for the attachment of the ear-drum, and sends down a rugged process, half an inch long, from its lower margin. In the _Dasypodes_ and _Myrmecophagæ_, the tympanic bone soon becomes anchylosed with the other parts of the temporal; it is only in _Orycteropus_, among the existing insectivorous _Bruta_ or _Edentata_, that it manifests throughout life the fœtal condition of a distinct bony hoop, deficient at the upper part. The os tympanicum of _Orycteropus_, however, differs from that of _Glossotherium_, in forming part of the circumference of an ellipse, whose long axis is vertical; and in sending outwards, from its anterior part, a convex eminence, which terminates in a point directed downwards and forwards.
Such appear to be the most characteristic features of the cranial fragment under consideration, in which we have found, that the articular surface for the os hyoides throws more light upon the nature of the animal of which it is a part, than even the glenoid cavity itself. There now remains to be described as much of the individual characters of the constituent bones as the specimen exhibits.
The occipital bone, besides forming the posterior and part of the inferior parietes of the cranium, extends for about half an inch upon the sides, where the ex-occipital element is articulated by a vertical suture with the mastoid element of the temporal: this suture is situated in a deep and well-marked muscular depression (_e_, fig. 1), measuring three inches in the vertical, and upwards of one inch in the transverse direction. The other sutures, uniting the occipital to the adjoining bones, are obliterated. The breadth of the occipital region must have exceeded the height of the same by about one-third. The condyle extends nearly to the external boundary of the occipital aspect of the cranium; there is situated, external to it, only a small ovate, rounded and smooth protuberance. The slightly concave surface of the occipital plane of the cranium is bounded above by a thick obtuse ridge, the muscular impressions are well sculptured upon it. It is traversed transversely at its upper third by a slightly elevated bony crest; and the surface below this ridge is again divided by a narrower intermuscular crest, which runs nearly vertically, at about an inch and a half from the external boundary of the occipital plane. As a similar crest must have existed on the opposite side, the general character of the occipital surface in the Glossothere would resemble that of the Toxodon. A similar correspondence may be noticed in the terminal position of the condyle, and the slope of the occipital plane.
Above the transverse ridge, the rough surface of the occipital plane slopes forward, at a less obtuse angle with the basal plane, to the first named ridge which separates the occipital from the coronal or superior surface of the skull. The contour of this surface runs forwards, as far as the fragment extends, in an almost straight line: the extent of surface between the temporal muscular ridges must have been about five inches posteriorly, but it decreases gradually as it extends forwards: all that part which is preserved is quite smooth. The attachment of the fasciculi of the temporal muscle, and the convergence of their fibres as they passed through the zygoma are well-marked on the sculptured surface of the bone. The zygomatic process is relatively stouter than in _Orycteropus_: it is prismatic: the external facet is nearly plane: the superior is concave, and increases in breadth anteriorly: the inferior surface offers a slight convexity behind the flattened articular surface for the lower jaw. The margin of the zygoma formed by the meeting of the upper and lower facets presents a semicircular curve, extended transversely from the cranium, and directed forwards.
The anterior extremity is obliquely truncated from below upwards and forwards, and presents a flattened triangular surface indicative of its junction with an os malæ: the space between this extremity and the side of the cranium measures one inch and nine lines across, and thus gives us the thickness of the temporal muscle. The distance from the origin of the zygoma to the occipital plane is relatively greater than in _Orycteropus_; _Glossotherium_ is in this respect more similar to _Myrmecophaga_ and _Manis_.
The sphenoid bone forms a somewhat smooth protuberance below and behind the base of the _zygoma_. The tympanic bone is wedged in between this protuberance in front, and the mastoid process behind. The chief peculiarity of the broad mastoid is the regular semicircular cavity at its under part for the articulation of the styloid bone of the tongue. This depression is separated below by a broad rough protuberance from the foramen jugulare, (_f_, fig. 2, Pl. XVI,) which is immediately external to, and slightly in advance of the great foramen condyloideum, _c_. A small rugged portion of the os petrosum separates the jugular from the carotid canal, which arches upwards and directly inwards to the side of the shallow sella turcica, (the external and internal orifices of the carotid canal are shown at _g_, figs. 2 and 3). The chief protuberance on the basis cranii is a large and rugged one, serving for the attachment of muscles, and due chiefly to the expansion of a great sinus in the body of the sphenoid. This protuberance is separated from the smaller sphenoid protuberance before mentioned by a large groove continued downwards and forwards from the tympanic cavity, and containing the Eustachian tube, which does not traverse a complete osseous canal. Immediately internal to the glenoid cavity is the large orifice of the canal transmitting the third division of the fifth pair of nerves, the principal branch of which endows the tongue with sensibility; this foramen (_h_, fig. 2) is rather less than that for the muscular nerve of the tongue.
The internal surface of the present cranial fragment affords a very satisfactory idea of the size and shape of the brain of the extinct species to which it belongs. It is evident that, as in other Bruta, the cerebellum must have been almost entirely exposed behind the cerebrum; and that the latter was of small relative size, not exceeding that of the Ass; and chiefly remarkable, as in the Orycterope, Ant-eater, and Armadillo for the great development of the olfactory ganglia. The antero-posterior extent of the cribriform plate, as exposed in this fragment, is three inches, and the complication of the œthmoid olfactory lamellæ which radiate from it into the nasal cavity is equal to that which exists in the smaller Edentata (fig. 3, Pl. XVI). The nasal cavity is complicated in _Glossotherium_ by the great number and capacious size of the air-cells which are in communication with it: these extend over all the upper, lateral, and back parts of the cranial cavity, as far even as the upper boundary of the foramen magnum: they also occupy the anterior two-thirds of the basis cranii. The external configuration of the skull would, therefore, afford a very inadequate or rather deceptive notion of the capacity of the cerebral cavity, were not the existence and magnitude of these sinuses known. The interspace of the outer and inner tables of the cranium are separated above the origins of the olfactory ganglia for the extent of three inches: above the middle of the cerebrum they are an inch and a half apart; at the sides of the cranium the interposed air-cells are from one to two inches across; at the back part of the cranium about one inch. The sinuses have generally a rounded form.
The foramen rotundum, (through which in figure 3 a probe is represented as passing), and the foramen ovale are situated close together, within a common transversely oblong depression (_i_). The carotid canal (_g_) opens into the outer side of the commencement of this wide channel, which conducts the great fifth pair of nerves to the outlets of its two chief divisions.
The petrous bone projects into the cranial cavity, in the form of an angular process with three facets: the foramen auditorium internum (_k_), and the aqueductus vestibuli, are situated on the posterior facet. Immediately behind the os petrosum is the foramen lacerum jugulare (_l_), situated at the point of convergence of the vertical groove of the lateral sinus, with a groove of similar size continued forwards from above the anterior condyloid canal. The plane of the internal opening of this canal (_c_, fig. 3) is directed obliquely inwards and backwards, and the lateral wall of the foramen magnum behind the foramen condyloideum slopes outwards to the edge of the condyle. Immediately internal to the foramen condyloideum is a small vascular foramen conducting a branch of the basilar artery into the condyloid canal, for the nourishment, doubtless, of the great lingual nerve.
In the relations of the plane of the internal orifice of the anterior condyloid foramen with that of the foramen magnum, we search in vain for a corresponding structure in any of the Mammiferous orders, save the Edentata:[30] and among these the Orycterope comes nearest the Glossothere in this respect. In the degree of development of the internal osseous ridge giving attachment to the tentorium cerebelli, the Ant-eaters and Armadillos more resemble the Glossothere than does the Orycterope; in which a continuous bony plate arches across the cranial cavity: in the Manis a still greater proportion of the tentorium is ossified, and it consequently recedes the furthest amongst the Edentata, in this, as in most other particulars of the cranial organization, from the Glossothere. The chief distinctive peculiarity in the cranium of the Glossothere, so far as it can be studied in the present fragment, and compared with that of other Edentata, is the deep, well-marked, semicircular styloid depression, above described.
A question may arise after perusing the preceding evidence, upon which the present fossil is referred to a great Edentate species nearly allied to the _Orycteropus_, whether one or other of the lower jaws, subsequently to be described, and in like manner referable, from their dentition, either to the _Orycteropodoid_ or _Dasypodoid_ families of Edentata, may not have belonged to the same species as does the present mutilated cranium. I can only answer, that those jaws were discovered by Mr. Darwin in a different and very remote locality,—that no fragments or teeth referable to them were found associated with the present fossil; and that, as it would be, therefore, impossible to determine from the evidence we have now before us, which of the two lower jaws should be associated with _Glossotherium_; and as both may with equal if not greater probability belong to a totally distinct genus, it appears to me to be preferable, both in regard to the advancement of our knowledge of these most interesting Edentata of an ancient world, as well as for the convenience of their description, to assign to them, for the present, distinct generic appellations.
The figures in Plate XVI. preclude the necessity of a table of admeasurements of the cranial fragment of _Glossotherium_.
DESCRIPTION OF A MUTILATED LOWER JAW AND TEETH, ON WHICH IS FOUNDED A
SUBGENUS OF MEGATHERIOID EDENTATA, UNDER THE NAME OF
MYLODON.
The genus _Megalonyx_, as is well known, owes its name and the discovery of the fossil remains on which it was founded, to the celebrated Jefferson,[31] formerly President of the United States. Cuvier, from an examination of a single tooth, and the casts of certain bones of the extremities, especially the terminal ones, determined the ordinal affinities of this remarkable extinct quadruped.[32] But while he retained, the name of _Megalonyx_, and used it in a generic sense, Cuvier offered no characters whereby other fossil remains might be generically either distinguished from, or identified with the _Megalonyx Jeffersonii_, unless, among such remains there happened to be a tooth, or a claw exactly corresponding with the descriptions and figures in the _Ossemens Fossiles_; and when, of course, a specific identity, and not merely a generic relationship would be established.
The greater part of Cuvier’s chapter on _Megalonyx_ is devoted to the beautiful and justly celebrated reasoning on the ungueal phalanx, whereby it is proved to belong, not to a gigantic Carnivore of the Lion-kind, as Jefferson supposed, but to the less formidable order of Edentate quadrupeds; and Cuvier, in reference to the tooth,—the part on which alone a generic character could have been founded,—merely observes that it resembles at least as much the teeth of one of the great Armadillos, as it does those of the Sloths.[33]
In the last edition of the _Rêgne Animal_, Cuvier introduces the _Megatherium_ and _Megalonyx_, between the Sloths and Armadillos; but alludes to no other difference between the two genera than that of size,—“l’autre, le _Megalonyx_, est un peu moindre.” (p. 226.) Some systematic naturalists, as Desmarest, and Fischer, have, therefore, suppressed the genus, and made the _Megalonyx_ a species of _Megatherium_ under the name of _Megatherium Jeffersonii_. The dental characters of the genus _Megatherium_ are laid down by Fischer[34] as follows:—“_Dent. prim, et lan. ⁰⁄₀., molar es ⁴⁄₄⁴⁄₄, obducti, tritores, coronide nunc planâ transversim sulcatâ nunc medio excavatâ marginibus prominulis._” That _Megalonyx_ had the same number of molares as _Megatherium_, (supposing that number in the Megathere to be correctly stated, which it is not,) is here assumed from analogy, for neither Jefferson, Wistar, nor Cuvier,—the authorities for _Megalonyx_ quoted by Fischer—possessed other means of knowing the dentition of that animal than were afforded by the fragment of a single tooth.
Now the almost entire lower jaw about to be described offers, in so far as respects the general form and structure of the teeth, the same kind and degree of correspondence with the _Megatherium_, as does the _Megalonyx Jeffersonii_ of Cuvier: and, what is only probable in that species, is here certain, viz., an agreement with the Megatherium in the class, viz. _molares_, to which the teeth exclusively belong. The question, therefore, on which I find myself, in the outset, called upon to come to a decision is, as to the preference of the mode of viewing the subject of the generic relationship of the _Megalonyx_ adopted by Desmarest, Fischer, &c., or of that, on which Cuvier, and after him Dr. Harlan, have practically acted: whether, in short, the genus _Megatherium_ is to rest upon the more comprehensive characters of kind and general structure of the teeth, or upon the more restricted ones, of form and such modifications in the disposition and proportions of the component textures of the tooth, as give rise to the characteristic appearances of the triturating surface of the crown.
With respect to existing Mammalia, most naturalists of the present day seem to be unanimous as to the convenience at least of founding a generic or subgeneric distinction on well-marked modifications in the form and structure of the teeth, although they may correspond in number and kind, in proof of which it needs only to peruse the pages of a _Systema Mammalium_ which relate to the distribution of the Rodent Order. According to this mode of viewing the logical abstractions under which species are grouped together, the extinct Edentate Mammal discovered by Jefferson must be referred to a genus distinct from _Megatherium_, and for which the term _Megalonyx_ should be retained. This will be sufficiently evident by comparing the descriptions given by Cuvier of one of the teeth of the _Megalonyx Jeffersonii_, and by Dr. Harlan of a tooth of his _Megalonyx laqueatus_, with those of the _Megatherium_ which have been published by Mr. Clift. The fragment of the molar tooth of the _Megalonyx Jeffersonii_, described and figured in the _Ossemens Fossiles_, seems to have been implanted in the jaw, like the teeth of the _Megatherium_, by a simple hollow base similar in form and size to the protruded crown: its structure Cuvier describes as consisting of a central cylinder of bone enveloped in a sheath of enamel.[35] The transverse section of this tooth presents an irregular elliptical form, the external contour being gently and uniformly convex, the internal one, undulating; convex in the middle, and slightly concave on each side, arising from the tooth being traversed longitudinally on its inner side by two wide and shallow depressions.
The imperfect tooth of the species called by Dr. Harlan _Megalonyx laqueatus_, and of which a cast was presented by that able and industrious naturalist to the Museum of the Royal College of Surgeons, resembles in general form, and especially in the characteristic double longitudinal groove on the inner side, the tooth of the _Megalonyx Jeffersonii_. It is thus described by Dr. Harlan:
“The fractured molar tooth appears to have belonged to the inferior maxilla on the right side; the crown is destroyed; a part of the cavity of the root remains. The body is compressed transversely, and presents a double curvature, which renders its anterior and exterior aspects slightly convex; the posterior and interior gently concave; these surfaces are all uniform, with the exception of the interior or mesial aspect, which presents a longitudinal rib or ridge, one-half the thickness of the long diameter of the tooth; with a broad, not profound longitudinal groove or channel along each of its borders. It is from this resemblance to a portion of a fluted column, that the animal takes its specific appellation (_Meg^x. laqueatus_).
“The crown would resemble an irregular ellipsis widest at the anterior portion. The tooth consists of a central pillar of bone surrounded with enamel, the former of a dead white, the latter of a ferruginous brown colour: the transverse diameter is more than two-thirds less than its length, whilst that of _Meg^x. Jeffersonii_ is only one-third less—the antero-posterior diameter is one-half its length in the former, and two-thirds less in the latter. The proportions of this tooth are consequently totally at variance with that of its kindred species.” [Vide Pl. XII. fig. 7, 8, 9.][36]
Dr. Harlan describes also two claws of the fore-foot, a radius, humerus, scapula, one rib, an os calcis, a metacarpal bone, certain vertebræ, a femur, and tibia, of the same _Megalonyx_; these parts of the skeleton, together with the tooth, which so fortunately served to establish the generic relationship of the species with the _Megalonyx_ of Jefferson and Cuvier, were discovered in Big-bone-cave, Tennessee, United States.
Dr. Harlan does not enter into the question of the generic characters of _Megalonyx_, but it would seem that he felt them to rest not entirely on dental modifications, for he observes that “a minute examination of the tooth and knee-joint renders it not improbable, supposing the last named character to be peculiar to it, that if the whole frame should hereafter be discovered, it may even claim a generic distinction, in which case, either Aulaxodon, or PLEURODON, would not be an inappropriate name.”[37]
There can be no doubt, as it appears to me, with respect to a fossil jaw presenting teeth in the same number, and of the same general structure, as in the _Megatherium_, and with individual modifications of form, as well-marked as those which distinguish _Megatherium_ from _Megalonyx_, that the Palæontologist has no other choice than to refer it, either as Fischer has done with _Megalonyx_, to a distinct species of the genus _Megatherium_, or to regard it as the type of a subgenus distinct from both. With reference, however, to the _Pleurodon_ of Dr. Harlan, after a detailed comparison of the cast of the tooth on which that genus is mainly founded, with the descriptions and figures of the tooth of the _Megalonyx Jeffersonii_, in the “Ossemens Fossiles,” they seem to differ in so slight a degree as to warrant only a specific distinction, and this difference even, viewing the various proportions of the teeth in the same jaw of the _Megatherium_, is more satisfactorily established by the characters pointed out by Dr. Harlan in the form and proportions of the radius, than by those in the tooth itself.
The next notice of the _Megalonyx_ which I have consulted, in the hope of meeting with additional and more precise information as to its real generic characters, is an account given by the learned Professor Doellinger,[38] of some fossil bones, collected by the accomplished travellers Spix and Martius in the cave of Lassa Grande, near the Arrayal de Torracigos, in Brazil. In this collection, however, it unfortunately happens that there are no teeth, but only a few bones of the extremities, including some ungueal phalanges, which Professor Doellinger concludes, from their shape, the presence of an osseous sheath for the claw, and the form of their articulation, to belong, without doubt, to an animal of the Megatherioid kind, about the size of an Ox. He particularly states that they are not bones of an immature individual; but that they agree sufficiently with Cuvier’s descriptions and figures of the _Megalonyx_ to be referred to that species of animal (zu dieses thierart;) and he adds, what is certainly an interesting fact, that the fossils in question form the first of the kind that had been discovered out of North America.
Subsequently to the discovery of these bones, and of those of the _Megalonyx laqueatus_ above alluded to, the remains of another great Edentate animal were found in North America, and were deposited in the Lyceum at New York; among these is a portion of the lower jaw with the whole dental series of one side. It is thus described by Dr. Harlan.
“The fragment I am now about to describe is a portion of the dexter lower jaw of the Megalonyx, containing four molar teeth; three of the crowns of these teeth are perfect, that of the anterior one is imperfect. These teeth differ considerably from each other in shape, and increase in size from the front, the fourth and posterior tooth being double the size of the first, and more compressed laterally; it is also vertically concave on its external aspect, and vertically convex on its internal aspect; the interior or mesial surface is strongly fluted, and it has a deep longitudinal furrow on the dermal aspect, in which respect it differs from the tooth of the _M. laqueatus_ previously described by me, of which the dermal aspect is uniform, but to which, in all other respects, it has a close resemblance. I suppose it therefore probable, that this last may have belonged to the upper jaw. The three anterior molars differ in shape and markings: they are vertically grooved, or fluted, on their interior and posterior aspects, a transverse section presenting an irregular cube. The length of the crown of the posterior molar is two inches: the breadth about five-tenths of an inch: the length of the tooth is three inches and six-tenths. The diameter of the penultimate molar is eight-tenths by seven-tenths of an inch. The length of this fragment of the jaw-bone is eight inches and four-tenths; the height three inches and six-tenths: the length of the space occupied by the alveolar sockets five inches and eight-tenths. The crown of the tooth presents no protuberances, but resembles that of the Sloth; the roots are hollow.”[39]
This fossil is referred by Dr. Harlan to his _Megalonyx laqueatus_; but, pending the absence of other proof of the identity of species, in which, as may be seen by comparing fig. 2, with fig. 4, in Pl. XVII., the teeth differ widely in form, it would be obviously hazardous to adopt such an approximation on hypothetical grounds.[40] In order, however, to obtain more satisfactory evidence of the nature and amount of the difference between the _Megalonyx laqueatus_, and the allied animal represented by the above-described fragment of lower jaw, I wrote to my much respected friend M. LAURILLARD, requesting him to send me a sketch of the teeth in the cast of that lower jaw, which had been transmitted from New York to the Garden of Plants. With full confidence in the characteristic precision and accuracy of the drawing with which I have been obligingly favoured by M. Laurillard, I am disposed to regard the amount of difference recognizable in every tooth in the lower jaw in question (fig, 3 and 4,) as compared with the molar tooth either of _Megalonyx Jeffersonii_ (fig. 1,) or _Meg^x. laqueatus_ (fig. 2) to be such as to justify its generic separation from _Megalonyx_ on the same grounds as _Megalonyx_ is distinguished from _Megatherium_, and for the subgenus of Megatherioid Edentata, thus indicated, I would propose the name of MYLODON.[41] The species of which the fossil remains are described by Dr. Harlan may be dedicated to that indefatigable Naturalist who has contributed to natural science so much valuable information respecting the Zoology, both recent and fossil, of the North American continent. The fossil about to be described represents a second and smaller species of the same genus, and I propose to call it _Mylodon Darwinii_, in honour of its discoverer, of whose researches in the Southern division of the New World it forms one of many new and interesting fruits.
This fossil was discovered in a bed of partly consolidated gravel at the base of the cliff called Punta Alta, at Bahia Blanca in Northern Patagonia: it consists of the lower jaw with the series of teeth entire on both sides: but the extremity of the symphysis, the coronoid and condyloid processes, and the angular process of the left ramus, are wanting. The teeth are composed, as in _Bradypus_, _Megatherium_ and _Megalonyx_, of a central pillar of coarse ivory, immediately invested with a thin layer of fine and dense ivory, and the whole surrounded by a thick coating of cement.
In the fig. 5, Pl. XVII., the fine ivory is represented by the white striated concentric tract on the grinding surface of the teeth; it is of a yellowish-white colour in the fossil, and stands out, as an obtuse ridge, from that surface: both these conditions depend on the large proportion of the mineral to the animal constituent in this substance of the tooth. The external layer of the cement presents in the fossil the same yellowish-brown tint as the bone itself, which it so closely resembles, both in intimate structure and in chemical composition; the internal layer next the dense ivory is jet black, indicating the great proportion of animal matter originally present in this part. The central pillar of coarse ivory, which, from its more yielding texture, has been worn down into a hollow at the triturating surface of the tooth, also presents, as a consequence of the less proportion of the hardening phosphates, a darker brown colour than the external layer of the cement, or the bone itself.
The teeth are implanted in very deep sockets; about one-sixth only of the last molar projects above the alveolus; the proportion of the exposed part of the tooth increases as they are placed further forwards. The implanted part of each tooth is simple; preserving the same size and form as the projecting crown, and presenting a large conical cavity at the base, indicative of the original persistent pulp, and perpetual growth of these teeth.
The extent of the whole four alveoli is four inches, eight lines; the length of the jaw from the angle to the broken end of the symphysis is seventeen inches and a half;[42] from the figures it will be seen that only a small proportion of the anterior part of the jaw is lost, so that we may regard the dentigerous part of the jaw as being limited to about one-fourth of its entire length; the alveoli being nearly equidistant from the two extremities. The first and second teeth, counting backwards, are separated by an interspace of rather more than three lines; that between the second and third is one line less; the third and fourth are rather more than a line apart: from the oblique position, however, of the three hinder teeth the intervals between them appear in a side view, as in fig. 1, Pl. XIX., to be less than in reality, and the third and fourth teeth seem to touch each other.
Each tooth has a form and size peculiar to itself, and different from the rest, but corresponds of course with its fellow on the opposite side. The same may be observed, but in a less degree, in the teeth of the Megatherium itself; hence, it is obviously hazardous to found a generic distinction upon a single tooth, unless, as in the case of the _Glyptodon_,[43] the modification of form happens to be extremely well-marked. The whole series of teeth, or their sockets, at least of one of the jaws, should be known for the purpose of making a satisfactory comparison with the previously established Edentate genera.
The first molar in the present jaw is the smallest and simplest of the series: its transverse section is ellipsoid, or subovate, narrowest in front, and somewhat more convex on the outer than on the inner side: the long diameter of the ellipse is nine lines, the short or transverse diameter six lines: the length of the tooth may be about three inches, but I have not deemed it necessary to fracture the alveolus in order to ascertain precisely this point.
The second tooth presents in transverse section a more irregular and wider oval figure than the first: the line of the outer side is convex, but that of the inner side slightly concave, in consequence of the tooth being traversed longitudinally by a broad and shallow channel or impression; the longitudinal diameter of the transverse section is one inch; the transverse diameter at the widest part nine lines. There is a slight difference in the size of this tooth on the two sides of the jaw, the right one, from which the above dimensions are taken, being the largest.
The transverse section of the third tooth has a trapezoidal or rhomboidal form; the angles are rounded off; the posterior one is most produced; the anterior and posterior surfaces are flattened, the latter slightly concave in the middle; the external and internal sides are concave in the middle, especially the inner side, where the concavity approaches to the form of an entering notch. The longest diameter of the transverse section of this tooth is thirteen lines, the shortest seven lines and a half: in the tooth on the right side the external surface is nearly flat; this slight difference is not indicated in the figure (Pl. XVIII.)
The last molar, which is generally the most characteristic in the fossil _Bruta_, presents in an exaggerated degree the peculiarities of the preceding tooth; the longitudinal channels on both the outer and inner surfaces encroach so far upon the substance of the tooth, that the central coarse ivory substance is as it were squeezed out of the interspace, and the elevated ridge of the dense ivory describes an hour-glass figure upon the triturating surface, the connecting isthmus being but half the breadth of the rest of the tract; the external cæmentum preserves nearly an equal thickness throughout. Of the two lobes into which this tooth is divided by the transverse constriction, the anterior is the largest; their proportions and oblique position are pretty accurately given in the figure. The longitudinal diameter of the transverse section of this tooth is one inch, seven lines, its greatest lateral or transverse diameter is ten lines, its least diameter at the constricted part is three lines, the length of the entire tooth is four inches. Judging from the form of the jaw, the length of the other teeth decreases in a regular ratio to the anterior one. The posterior tooth is slightly curved, as shown in fig. 2, Pl. XIX., with the concavity directed towards the outer side of the jaw.
The general form of the horizontal ramus of the jaw, is so well illustrated in the figures Pl. XVIII. and XIX., that the description may be brief.
The symphysis is completely anchylosed, about four inches in length, and extended forward to the extremity of the jaw at a very slight angle with the inferior border of the ramus: it is of great breadth, smooth and gently concave internally, and suggests the idea of its adaptation for the support and gliding movements forwards and backwards of the free extremity of a long and well-developed tongue.
The exterior surface of the symphysis is characterized by the presence of two oval mammilloid processes, situated on each side of the middle line, and about half-way between the anterior and posterior extremes of the symphysis. A front view of these processes, of the natural size, is given in fig. 4, Pl. XIX.: a side view of the one on the right side represented in the reduced figure.
Nearly four inches behind the anterior extremity of the above process is the large anterior opening of the dental canal: it is five lines in diameter, situated about one-third of the depth of the ramus of the jaw from the upper margin. The magnitude of this foramen, which gives passage to the nerve and artery of the lower lip, indicates that this part was of large size; and the two symphyseal processes, which probably were subservient to the attachment of large retractor muscles, denote the free and extensive motions of such a lip, as we have presumed to have existed from the size of the foramina destined for the transmission of its nervous and nutrient organs.
The angle of the jaw is produced backwards, and ends in an obtuse point, slightly bent upwards; a foramen, one-third less than the anterior one, leads from near the commencement of the dental canal, to the outer surface of the jaw, a little below and behind the last molar tooth; this foramen presents the same size and relative position on both sides of the jaw. I find no indication of a corresponding foramen, or of symphyseal processes in the figures or descriptions of the lower jaw of the Megatherium, nor in the lower jaw of the Sloths, Ant-eaters, Armadillos, or Manises, which I have had the opportunity of examining with a view to this comparison.
In the Megatherium the inferior contour of the lower jaw is peculiarly remarkable, as Cuvier has observed, for the convex prominence or enlargement which is developed downwards from its middle part. In the Mylodon the corresponding convexity exists in a very slight degree, not exceeding that which may be observed at the corresponding part of the lower jaw of the Ai, or Orycterope. A broad and shallow furrow extends along the outer side of the jaw, close to the alveolar margin, from the beginning of the coronoid process to the anterior dental foramen.
The base of the coronoid process begins external and posterior to the last grinder: the whole of the ascending ramus of the jaw, beneath the coronoid process is excavated on its inner side by a wide and deep concavity, bounded below by a well-marked ridge, which extends obliquely backwards from the posterior part of the alveolus of the last grinder to the inferior margin of the ascending ramus, which is bent inwards before it reaches the angle of the jaw.
The large foramen or entry to the dental canal is situated in the internal concavity of the ascending ramus of the jaw, two inches behind the last molar, three inches from the lower margin of the ramus, and nearly five inches from the elevated angle of the jaw: it measures nine lines in the vertical diameter, and its magnitude indicates the large size of the vessels which are destined to supply the materials for the constant renewal of the dental substance,—a substance which from its texture must be supposed to have been subject to rapid abrasion. About an inch behind the dental foramen a deep vascular groove, about two lines in breadth, is continued downwards to the ridge which circumscribes the internal concavity of this part of the jaw, and perforates the ridge, which thus arches over the canal: this structure is present in both rami of the jaw. The mylo-hyoid ridge is distinctly marked about an inch and a half below the alveolar margin. Other muscular ridges and irregular eminences are present on the outer side of the base of the ascending ramus, and near the angle of the jaw; as shown in fig. 1, Pl. XIX.
From the preceding descriptions it will be seen that the lower jaw of the _Mylodon_ is very different from that of the _Megatherium_; with that of the _Megalonyx_ we have at present no means of comparing it. Among existing Edentata the Mylodon, in the form of the posterior part and angle of the jaw, holds an intermediate place between the Ai and the great Armadillo; in the form of the anchylosed symphysis of the lower jaw it resembles most closely the Unau or two-toed Sloth; but in the peculiar external configuration of the symphysis resulting from the mammilloid processes above described, the Mylodon presents a character which has not hitherto been observed in any other species of _Bruta_, either recent or fossil.
In conclusion it may be stated, that the teeth and bones here described offer all the conditions and appearances of those of a full grown animal; and that they present a marked difference of size as compared with those of the _Mylodon Harlani_, as will be evident by the following admeasurements.
ADMEASUREMENTS OF THE LOWER JAW OF MYLODON DARWINII.
Inches. Lines. Length (as far as complete) 17 6 Extreme width, from the outside of one ramus to that of the other 9 0 Depth of each ramus 4 9 Length of alveolar series 4 8 From first molar to broken end of symphysis 6 0 Breadth of symphysis 3 7 Longitudinal extent of symphysis 4 6 Circumference of narrowest part of each ramus 5 9
DESCRIPTION OF A CONSIDERABLE PART OF THE SKELETON OF A LARGE EDENTATE
MAMMAL, ALLIED TO THE MEGATHERIUM AND ORYCTEROPUS, AND FOR WHICH IS
PROPOSED THE NAME OF
SCELIDOTHERIUM[44] LEPTOCEPHALUM.
Of the large Edentate quadrupeds that once existed in the New World, sufficient of the osseous remains of the gigantic Megatherium alone has been transmitted to Europe to give a satisfactory idea of the general form and proportions of the extinct animal.
Different bones of the Megalonyx, Mylodon, and Glyptodon have been described, but not sufficient of the remains of any individual of these subgenera has, hitherto, reached Europe, or been so described as to enable us to form a comparison between them and the Megatherium, or any of the existing Edentata, in regard to the general construction and proportions of the entire skeleton.
This state of our knowledge of the osteology of the singular giants of the Edentate Order renders the remains of the present animal peculiarly interesting, since, although the extremities are too imperfect to enable us to reconstruct the entire skeleton, a sufficient proportion of it has been preserved in the natural position to give a very satisfactory idea of its affinities to other Edentata, whose osteology is more completely known.
The fossil remains here described were discovered by Mr. Darwin in the same bed of partly consolidated gravel at Punta Alta, Northern Patagonia, as that in which the lower jaws of the _Toxodon_ and _Mylodon_ were imbedded. The parts of the skeleton about to be described were discovered in their natural relative position, as represented at Pl. XX., indicating, Mr. Darwin observes, that the sublittoral formation in which they had been originally deposited had been subject to little disturbance.[45] They include the cranium, nearly entire, with the teeth and part of the os hyoides; the seven cervical, eight of the dorsal, and five of the sacral vertebræ, the two scapulæ, left humerus, radius and ulna, two carpal bones, and an ungueal phalanx; both femora, the proximal extremities of the left tibia and fibula, and the left astragalus.
The principal parts of the cranium which are deficient are the anterior extremities of both the upper and lower jaws, the os frontis, æthmoid bone, and the whole upper part of the facial division of the skull; but sufficient remains to show that the general form of the skull resembled an elongated, slender, subcompressed cone, commencing behind by a flattened vertical base, slightly expanding to the zygomatic region, and thence gradually contracting in all its dimensions to the anterior extremity.
The Cape Ant-eater (_Orycteropus_), of all Edentata, most nearly resembles the present fossil in the form of its cranium, and next in this comparison the great Armadillo (_Dasypus gigas_, Cuv.) may be cited: on the supposition, therefore, that the correspondence with the above existing Edentals observable in the parts of the fossil cranium which do exist, was carried out through those which are defective, the length of the skull of the Scelidothere must have been not less than two feet. If now the reader will turn to Pl. XX. he will see that this cranium is singularly small and slender in proportion to the rest of the skeleton, especially the bulky pelvis and femur, of which bones the latter has a length of seventeen inches, and a breadth of not less than nine inches; the astragalus, again, exceeds in bulk that of the largest Hippopotamus or Rhinoceros; yet the condition of the epiphyseal extremities of the long bones proves the present fossils to have belonged to an immature animal. Hence, although the Scelidothere, like most other Edentals, was of low stature, and, like the Megatherium, presented a disproportionate development of the hinder parts, it is probable, that, bulk for bulk, it equalled, when alive, the largest existing pachyderms, not proboscidian. There is no evidence that it possessed a tesselated osseous coat of mail.
I shall commence the description of the present skeleton with the cranium. The condyles of the occiput (See Pl. XXI. fig. 2,) are wide apart, sub-elliptic, very similar in position, form, and relative size to those in _Orycteropus_. The foramen occipitale is transversely oval, its plane slopes from above downwards and forwards at an angle of 40° with that of the occipital region of the skull. This region, as before stated, is vertical in position (see fig. 1, Pl. XXI.), of a sub-semicircular form, the breadth being nearly one-third more than the height; it is bounded above and laterally by a pretty regular curve; but the superior margin is not produced so far backwards as in _Orycteropus_. The occipital plane is bisected by a mesial vertical ridge; there is a less developed transverse curved intermuscular crest which runs parallel with and about half an inch below the marginal ridge: the surface of the occipital plane on the interspaces of these ridges is irregularly pitted with the impression of the insertion of powerful muscles. The corresponding surface is smooth in the Orycterope and Armadillos; in the great extinct Glossothere it resembles in character that of the Scelidothere; but in the forward slope of the occipital plane the Glossothere differs in a marked degree from the present animal.
The upper surface of the cranium is smooth and regularly convex. The extent of the origin of the temporal muscles is defined by a slightly-raised broad commencement of a ridge, which, in the older animal, might become more developed. There is no trace of this ridge in the Orycterope; but it exists in the Armadillos, in which the teeth are of a denser texture, and better organized for mastication, and consequently are associated with better developed masticatory muscles. It will be subsequently shown that the Scelidothere resembles the Armadillos in so far as it possesses a greater proportion of the dense ivory to the external cæmentum in its teeth, than does the Megatherium; while it differs widely from the Orycterope, in the structure of its teeth. The teeth, however, are fewer in the Scelidothere than in any Armadillo, and relatively smaller than in most of the species of that family. Accordingly we find that the zygomatic arches are relatively weaker; and in this particular the _Scelidothere_ corresponds with the Orycterope. The zygomatic process of the temporal commences posteriorly about an inch and a half from the occipital plane, its origin or base is extended forwards in a horizontal line fully four inches, where it terminates as usual in a thin concave edge, as shown on the right side in Pl. XXII. The free portion of the zygoma, continued forwards from the outer part of this edge, is a slender sub compressed process, half an inch in the longest or vertical diameter, and less than three lines in the transverse; the extremity of this process is broken off; the opposite extremity of the malar portion of the zygoma is entire, and obtusely rounded. The bony arch may have been completed by the extension of the temporal process to the malar one, but the two parts undoubtedly were not connected together by so extensive a surface as in the Orycterope. On the other hand, if the zygomatic arch be naturally incomplete in the Scelidothere, the interspace between the malar and temporal portions must be relatively much less than in the Sloth or Ant-eater; for the broken end of the temporal part is separated from the obtusely rounded apex of the malar process in the present specimen by an interval of only one inch.
The articular surface (Pl. XXIII., fig. 2) beneath the zygoma for the lower jaw is flat and even, with the outer and inner margin slightly bent down, but having no definable anterior or posterior limits; its breadth is two inches. It differs from the corresponding surface in the Orycterope in being separated by a relatively wider interval from the tympanic bone, and in wanting consequently the support which the bony meatus auditorius gives in the Orycterope to the back part of the mandibular joint. The Armadillos differ still more from the Scelidothere in this important part of the cranial organization, inasmuch as the glenoid cavity is not only protected behind by the descending os tympanicum, but also in front by a corresponding vertical downward extension of the os malæ. The Scelidothere in the general form and relative position of the surface for the articulation of the lower jaw resembles the Glossothere more closely than any other Edentate animal with which I have been able to compare it.
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The zoology of the voyage of H.M.S. Beagle [vol. 1 of 5]Chapter IV: Part 4
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