Part 13

The fragments of the dorsal vertebræ and ribs of the Scelidotherium, which are figured in Plate XX., offer no modifications which need detain our attention; they closely conform, excepting in the greater relative height of the anterior dorsal spines, already noticed, with the Megatherioid type. The sacrum manifests in its vast expanse, the great development of the posterior transverse processes to join the ischium, the capacious medullary cavity, and wide nervous foramina, a like conformity with the Megatherium, and a corresponding harmony with the disproportionate bulk of the hind legs.

BONES OF THE EXTREMITIES.

The Scapula in its double spine, the osseous arch formed by the confluence of the acromion with the coracoid process, and the substitution of a distinct foramen for the suprascapular notch, agrees with that of the Megatherium: but the span of the acromial arch is relatively wider, and the surface for the articulation of the clavicle is better marked. This articular surface, which is distinctly shewn upon the acromion of both the scapulæ in Pl. XX. is the more interesting, as being the only evidence of the clavicle of the Scelidothere which we at present possess; but it is enough to prove that this quadruped enjoyed all the advantages in the actions of the fore extremity, which arise out of the additional fixation of the shoulder-joint afforded by the clavicle—a bone which the extinct Megatherioids are the largest of the mammiferous class to possess in a completely developed state. The form, position, and aspect of the glenoid cavity for the humerus closely correspond with the condition of the same part in the Megatherium. The limits of the acromial and coronoid portions of the arch were still defineable in the present skeleton, which indicates the nonage of the individual in the unanchylosed condition of most of the epiphysiaes.

In regard to the presence of a clavicle in the Megalonyx M. Lund has deduced certain conclusions, which, if well founded, would be equally applicable to the present allied species, and to the great Megatherium. I am induced, therefore, to offer a few physiological observations on that bone, which appear to me to lead to a more correct interpretation of its uses and relations in the great mammiferous animals now under consideration.

When the anterior extremities in mammalia are used simply for the purpose of progressive motion on dry land, as in the Pachyderms and Ruminants, or in water, as in the Cetaceans, there is no clavicle; this bone is introduced between the sternum and acromion, in order to give firmness and fixity to the shoulder-joint when the fore-leg is to discharge some other office than that of locomotion. In these cases, however, the clavicle exists in various degrees of development, and even its rudiment may be dispensed with in some of the actions which require a considerable extent of lateral or outward motion, and of freedom of rotation of the fore-limb. When, therefore, we find the clavicle fully developed in the skeleton of an extinct mammiferous animal, and so placed as to give the humeral articulation all the benefit of this additional mechanism, we may confidently expect that it will afford an insight into the habits and mode of life of such extinct species. M. Lund has argued from the clavicle of the Megalonyx, that it climbed like a Sloth. “Animals,” says Sir C. Bell, “which fly or dig, or climb, as Bats, Moles, Porcupines, Squirrels, Ant-eaters, Armadillos, and Sloths, have this bone; for in them, a lateral or outward motion is required.” But in regard to the present problem, we have to enquire whether the clavicle manifests any modifications of form, of strength, or development in relation to the special differences of these several actions, with which its presence is asserted to be associated?

In mammals which fly, the clavicle is always complete: the rabbit, the fox, and the badger are instances of burrowing animals in which the clavicle is absent or rudimental. The presence of a perfect clavicle is not more constant in climbing quadrupeds. The Ai, for example, has an incomplete clavicle, which is attached to the acromion process, and terminates in a point about one-fourth of the distance between the acromion and the top of the sternum, to which the clavicular style is attached by a long slender ligament: the advantage, therefore, which a perfect clavicle affords in the fixation of the shoulder-joint, is lost to this climber par excellence. Again, the Bears, which are the bulkiest quadrupeds that are gifted with the faculty of climbing, and this in so perfect a degree that the Sun-bears of the Eastern Tropics may be termed arboreal animals,—these scansorial quadrupeds are destitute of even the smallest rudiment of a clavicle, as I have ascertained by repeated careful dissection.

Since, therefore, a clavicle in any degree of development is not essential to a climbing quadruped, we must seek for some other relation and use of that remarkably strong, and perfect bone, as it exists in the Megathere, Megalonyx, and Scelidothere. The absence of ‘dentes primores’ or of anterior or incisive teeth in these quadrupeds at once sets aside any idea of its connection with an action of the fore extremities, very common in the mammals which possess clavicles, viz., that of carrying the food to the mouth, and holding it there to be gnawed by the teeth. Flying is of course out of the question, although our surprise would hardly be less at seeing a beast as bulky as an elephant climbing a tree, than it would be to witness it moving through the air. If now we restrict our comparison to the relations of the clavicle in that order of Mammalia to which the extinct species in question belonged, we shall see that it is most constant, strongest, and most complete in those species which make most use of their strong and long claws in displacing the earth, as the Armadillos and Orycteropus: and, as the clavicle is incomplete in one climbing Edental, we are naturally led to conclude that its perfect development in an extinct species must have been associated with uses and relations analogous to those with which it coexists in other genera of the same order. Thus it will be seen, that, in rejecting the conclusion drawn by M. Lund from the presence of a clavicle, I concur in the opinion expressed by Dr. Buckland that the Megatherium—and with it the Megalonyx and Scelidotherium—had the shoulder-joint strengthened by the clavicle, in reference to the office of the fore-arm, as an instrument to be employed in digging roots out of the ground. Not, however, that these gigantic quadrupeds fed on roots, but rather, as the structure of the teeth would show, on the foliage of the trees uprooted by the agency of this powerful mechanism of the fore-legs, and of the otherwise unintelligible colossal strength of the haunches, hind legs, and tail.

The humerus presents a large convex oval head, on each side of which is a tuberosity for the implantation of the supra- and sub-scapular muscles: these tuberosities do not rise above the articular convexity, so as to restrict the movements of the shoulder-joint, as in the Horse and Ruminants, but exhibit a structure and disposition conformable to those which characterize the proximal extremity of the humerus in other mammalia which enjoy rotatory movements of the upper or fore-limb. The tuberosities are, however, relatively more developed, and give greater breadth to the proximal end of the humerus in the Scelidothere than in the Megathere. The distal end of the humerus, although mutilated, clearly indicates that it had the same characteristic breadth of the external and internal condyles, as in the Megatherium. In fig. 1. Pl. XXV. which gives a front view of the left humerus, the broad internal condyle, with its extremity broken off, is seen projecting to the left hand; both in this figure and in fig. 2. in which the internal side of the humerus is turned towards the observer, the wide groove, with its two osseous boundaries, is shewn, which plainly indicates that the left condyle was perforated for the direct passage of the artery or median nerve, or of both, to the fore-arm. The groove for the musculo-spiral nerve on the outer side of the humerus is over-arched at its upper part by a strong obtuse process; which is comparatively less developed in the Megatherium. The trochlear or inferior articular surface of the humerus presents, as in the Megatherium, two well-marked convexities, with an intervening concavity: this indication of the rotatory power of the fore-leg is confirmed by the form of the head of the radius.

In Pl. XXV. fig. 4. a view is given of this articular surface: it presents the form of a subcircular gentle concavity, which plays upon the outer convexity of the humeral articular surface: immediately below the upper concavity the radius presents a lateral smooth convex surface, which rotates upon a small concavity on the ulna, analogous to the ‘lesser semilunar,’ in human anatomy, in which the mechanism for rotation, so far as the upper joint of the radius is concerned, is not more elaborately wrought out than in the present extinct edentate quadruped. The radius expands as it proceeds to the elbow-joint, where it attains a breadth indicative of the great power and size of the unguiculate paw, of which it may be called the stem, and to the movements of which it served as the pivot.

All the bones of the fore-limb just described—the scapula, the humerus, and the radius,—indicate by the bold features and projections of the muscular ridges and tubercles the prodigious force which was concentrated upon the actions of the fore-paw, and the ulna, in its broad and high olecranon (of which a side view is given in fig. 2. Pl. XXV.) gives corresponding evidence. The great semilunar concavity is traversed by a sub-median smooth ridge, which plays upon the interspace of the two humeral convexities. The body of the bone is subcompressed, straight, and diminishes in size as it approaches the carpal joint: the immediate articulating surfaces are wanting in both the radius and ulna, the epiphyseal distal extremities having become detached from their respective diaphyses.

Of the terminal segment of the locomotive extremities, the only evidence among the remains of the skeleton of the Scelidothere is the ungueal phalanx figured at Pl. XXVII. 3, 4, and 5; but as it is uncertain whether it belongs to the fore or hind-foot, it will be described after the other bones of the extremities have been noticed.

Of these bones the femur is the most remarkable, both for its great proportional size, and its extreme breadth, as compared with its length or thickness: but in all these circumstances the affinity of the Scelidothere with the Megathere is prominently brought into view. There is no other known quadruped with which the Scelidothere so closely corresponds in this respect. In proceeding, however, to compare together the thigh-bones of these two extinct quadrupeds, several differences present themselves, which are worthy of notice: of these the first is the presence in the Scelidothere of a depression for a ‘ligamentum teres’ on the back part of the head of the femur, near its junction with the neck of the bone: this is shewn in the posterior view of the femur given in Pl. XX. The head itself forms a pretty regular hemisphere: the great trochanter does not rise so high as in the Megatherium, but, relatively, it emulates it in breadth: the small trochanter is proportionally more developed: the external contour of the shaft of the femur is straighter in the Scelidothere than in the Megathere, and the shaft itself is less bowed forwards at that part. The articular condyles occupy a relatively smaller space upon the distal extremity of the femur in the Scelidothere, and they differ more strikingly from those of the Megathere, in being continued one into the other: the rotular surface, for example, which is shewn in fig. 5. Pl. XXV. is formed by both condyles, while in the Megatherium it is a continuation exclusively of the external articular surface.

The patella, which works upon the above-mentioned surface, is a thick strong ovate bone, with the smaller end downwards: rough and convex externally, smooth on the internal surface, which is concave in the vertical and convex in the transverse directions.

Of the bones of the leg only the proximal end of the tibia is preserved; but this is valuable, as shewing another well-marked difference between the Scelidothere and Megathere; for whereas in the latter the fibula is anchylosed with the tibia, this bone, in the Scelidothere, presents a smooth flat oval articular surface, which is shewn in fig. 2. Pl. XXVII. below the outer part of the head of the bone; from the size and appearance of which, I infer, that the fibula would not have become confluent with the tibia, even in the mature and full-grown animal.

The relative length of the fore and hind extremities cannot be precisely determined from the present imperfect skeleton of the Scelidothere; but there is good evidence for believing, that the fore extremity was the shortest. The humerus is shorter than the femur by one-ninth part of the latter bone; and the radius, which wants only the distal epiphysis, must have been shorter than the humerus. Now the relative development of the fore and hind legs is one of the points to be taken into consideration in an attempt to determine the habits and nature of an extinct mammal.

In climbing animals the prehensile power is more essential to the hinder than to the fore parts or extremities. In the leech the principal sucker is in the tail; and higher organized climbers, in like manner, depend mainly on their posterior claspers in descending trees, and hold on by means of them whilst selecting the place for the next application of those at the fore part of the body, whether their place be supplied by the beak, as in the Maccaws, or the fore-feet or hands in the Mammalia.

But, although we perceive the hinder limbs to be the last to lose the advantageous structure of the hand in the Quadrumanous species, and notwithstanding that the tail is for this purpose sometimes specially organized to serve as a prehensile instrument, yet we find that the power of grasping the branches of trees by either legs or tail is never maintained at the expense of undue bulk and weight of those organs. On the contrary, as the fore-limbs are the main instruments in the active exertions of climbing, so they are the strongest as well as the longest in all the best climbers, and the weight of the body which they have to drag along is diminished by dwarfish proportions of the hinder limbs, as in the Orangs and the Sloths.

Can those huge quadrupeds have been destined to climb that had the pelvis and hinder extremities more ponderous and bulky in proportion to the fore parts of the body than in any other known existing or extinct vertebrate animals?

M. Lund argues for the scansorial character of the Megalonyx, because its anterior extremities are longer than the posterior ones; but if they somewhat exceed the hind legs in length, how vastly inferior are they in respect of their breadth and thickness. The prehensile faculty of the hinder limbs of the best climbers, as the Sloths, Orangs, and Chameleons is by no means dependent on the superior mass of muscle and bone which enters into their conformation, but is associated with the very reverse conditions.

It is impossible to survey the discrepancy of size between the femur and the humerus of the Scelidothere, as exhibited in Pl. XX., without a conviction that it relates to other habits than those of climbing trees. The expanse of the sacrum, the evidence of the muscular masses employed in working the hind legs and tail, which is afforded by the capacity of the cavity lodging the part of the spinal marrow from which the nerves of those muscles were derived, both indicate the actions of the hind legs and tail to have been more powerful and energetic than would be required for mere prehension: and the association of hinder extremities so remarkable for their bulk, with a long and powerful tail, forbids my yielding assent to the speculation set forth by M. Lund, as to the prehensile character of the tail of the Megalonyx.

Astragalus.—In the examination of this characteristic bone I have kept in view the question of the habits of the Megatherioid quadrupeds in general, and the especial affinities of the Scelidotherium, in illustration of which I shall notice at the same time the peculiarities of the astragalus of the Sloth, Megatherium and Armadillo.

The upper articular surface of the astragalus of the Scelidotherium (Pl. XXVI. fig. 4.), presents, in its transverse contour, two convex pulleys, a and b, and an intermediate concavity, forming one continuous articular surface. The external or fibular trochlea (a) is strictly speaking convex only at its posterior part, the upper surface gradually narrowing to a ridge, as it advances forwards from which, the inner and outer parts slope away at an angle of 35°.

The tibial convexity (b) is more regular and less elevated, it has only half the antero-posterior extent of the outer pulley; its marginal contour forms an obtuse angle at the inner side.

In the Megatherium the upper articular surface of the astragalus is also divided into two trochleæ, of which the one on the fibular side (fig. 3, a), is of much greater relative size and extent than the tibial one (b), and is raised nearly four inches above the level of the latter, although in the oblique position in which the bone is naturally placed in the skeleton, the highest part of each convexity is on the same level. The fibular trochlea differs also from that in the Scelidothere in being regularly convex in the transverse as well as the antero-posterior direction. The tibial convexity resembles that in the Scelidothere, save in its smaller relative size; its internal margin likewise forms an angular projection below the internal malleolus.

The upper surface of the astragalus of the Mylodon, or Megalonyx(?) (Pl. XXVIII. fig. 5.), differs from that in the Megatherium in having a narrower fibular trochlear ridge.

The astragalus of the Ai (Bradypus tridactylus) differs widely from that of either the Megathere, Mylodon(?) or Scelidothere in having a conical cavity on the upper surface, in place of the fibular convexity, in which concavity the distal end of the fibula rotates like a pivot. This mechanism is closely related to the scansorial uses of the inwardly inflected foot of the Sloth.

If the astragalus of an Armadillo were placed side by side with that of the Megathere, it would be very difficult to determine the analogous parts, especially of the upper surface, unless guided by the intermediate structure presented by the Scelidothere. The upper surface of this bone, in the Armadillo, is, however, divided into two transversely convex trochleæ, separated by a much wider transversely concave surface. The fibular trochlea resembles that of the Scelidothere in having its upper and outer facets sloping away at an acute angle, but without meeting at a ridge anteriorly; this surface is not more raised above the tibial trochlea than in the Scelidothere.

The inner trochlea differs from that of the Scelidothere in having a greater relative antero-posterior extent, and in forming, in place of an uniform convex surface, a trochlea similar in structure to that on the outer side. The extent of rough surface on the upper part of the astragalus intervening between the articular surface for the bones of the leg, and that for the scaphoides is extremely small in the Megathere and Mylodon(?); it is relatively greater in the Scelidothere; it is still more extensive in the Armadillo; but is the longest in the Sloth. The anterior extremity of the astragalus which is entirely occupied by the scaphoid articular surface is very peculiar in the Scelidothere (Pl. XXVI. fig. 2.): it presents one convex and two concave facets, which, however, form part of one continuous articular surface: the convex facet forms the internal part of the surface, and presents a rhomboidal form with the long axis vertical. The concave facets (c and d) are extended transversely and placed one above the other; they are slightly concave in the transverse, and nearly flat in the vertical directions.