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 (Megx. 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 Megx. 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 Megx. 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 |