The Variation of Animals and Plants under Domestication

by Charles Darwin

The occipital foramen (fig. 10) presents some remarkable differences: in the wild rabbit, the lower edge between the condyles is considerably and almost angularly hollowed out, and the upper edge is deeply and squarely notched; hence the longitudinal axis exceeds the transverse axis. In the skulls of the lop-eared rabbits the transverse axis exceeds the longitudinal; for in none of these skulls was the lower edge between the condyles so deeply hollowed out; in five of them there was no upper square notch, in three there was a trace of the notch, and in two alone it was well developed. These differences in the shape of the foramen are remarkable, considering that it gives passage to so important a structure as the spinal marrow, though apparently the outline of the latter is not affected by the shape of the passage.

In all the skulls of the large lop-eared rabbits, the bony auditory meatus is conspicuously larger than in the wild rabbit. In a skull 4·3 inches in length, and which barely exceeded in breadth the skull of a wild rabbit (which was 3·15 inches in length), the longer diameter of the meatus was exactly twice as great. The orifice is more compressed, and its margin on the side nearest the skull stands up higher than the outer side. The whole meatus is directed more forwards. As in breeding lop-eared rabbits the length of the ears, and their consequent lopping and lying flat on the face, are the chief points of excellence, there can hardly be a doubt that the great change in the size, form, and direction of the bony meatus, relatively to this same part in the wild rabbit, is due to the continued selection of individuals having larger and larger ears. The influence of the external ear on the bony meatus is well shown in the skulls (I have examined three) of half-lops (see fig. 5), in which one ear stands upright, and the other and longer ear hangs down; for in these skulls there was a plain difference in the form and direction of the bony meatus on the two sides. But it is a much more interesting fact, that the changed direction and increased size of the bony meatus have slightly affected on the same side the structure of the whole skull. I here give a drawing (fig. 11) of the skull of a half-lop; and it may be observed that the suture between the parietal and frontal bones does not run strictly at right angles to the longitudinal axis of the skull; the left frontal bone projects beyond the right one; both the posterior and anterior margins of the left zygomatic arch on the side of the lopping ear stand a little in advance of the corresponding bones on the opposite side. Even the lower jaw is affected, and the condyles are not quite symmetrical, that on the left standing a little in advance of that on the right. This seems to me a remarkable case of correlation of growth. Who would have surmised that by keeping an animal during many generations under confinement, and so leading to the disuse of the muscles of the ears, and by continually selecting individuals with the longest and largest ears, he would thus indirectly have affected almost every suture in the skull and the form of the lower jaw!

Fig. 11—Skull of Half-lop Rabbit.

In the large lop-eared rabbits the only difference in the lower jaw, in comparison with that of the wild rabbit, is that the posterior margin of the ascending ramus is broader and more inflected. The teeth in neither jaw present any difference, except that the small incisors, beneath the large ones, are proportionately a little longer. The molar teeth have increased in size proportionately with the increased width of the skull, measured across the zygomatic arch, and not proportionally with its increased length. The inner line of the sockets of the molar teeth in the upper jaw of the wild rabbit forms a perfectly straight line; but in some of the largest skulls of the lop-eared this line was plainly bowed inwards. In one specimen there was an additional molar tooth on each side of the upper jaw, between the molars and premolars; but these two teeth did not correspond in size; and as no rodent has seven molars, this is merely a monstrosity, though a curious one.

The five other skulls of common domestic rabbits, some of which approach in size the above-described largest skulls, whilst the others exceed but little those of the wild rabbit, are only worth notice as presenting a perfect gradation in all the above-specified differences between the skulls of the largest lop-eared and wild rabbits. In all, however, the supra-orbital plates are rather larger, and in all the auditory meatus is larger, in conformity with the increased size of the external ears, than in the wild rabbit. The lower notch in the occipital foramen in some was not so deep as in the wild rabbit, but in all five skulls the upper notch was well developed.

The skull of the Angora rabbit, like the latter five skulls, is intermediate in general proportions, and in most other characters, between those of the largest lop-eared and wild rabbits. It presents only one singular character: though considerably longer than the skull of the wild rabbit, the breadth measured within the posterior supra-orbital fissures is nearly a third less than in the wild. The skulls of the silver-grey, and chinchilla and Himalayan rabbits are more elongated than in the wild, with broader supra-orbital plates, but differ little in any other respect, excepting that the upper and lower notches of the occipital foramen are not so deep or so well developed. The skull of the Moscow rabbit scarcely differs at all from that of the wild rabbit. In the Porto Santo feral rabbits the supra-orbital plates are generally narrower and more pointed than in our wild rabbits.

As some of the largest lop-eared rabbits of which I prepared skeletons were coloured almost like hares, and as these latter animals and rabbits have, as it is affirmed, been recently crossed in France, it might be thought that some of the above-described characters had been derived from a cross at a remote period with the hare. Consequently I examined skulls of the hare, but no light could thus be thrown on the peculiarities of the skulls of the larger rabbits. It is, however, an interesting fact, as illustrating the law that varieties of one species often assume the characters of other species of the same genus, that I found, on comparing the skulls of ten species of hares in the British Museum, that they differed from each other chiefly in the very same points in which domestic rabbits vary,—namely, in general proportions, in the form and size of the supra-orbital plates, in the form of the free end of the malar bone, and in the line of suture separating the occipital and frontal bones. Moreover two eminently variable characters in the domestic rabbit, namely, the outline of the occipital foramen and the shape of the “raised platform” of the occiput, were likewise variable in two instances in the same species of hare.

Vertebræ.—The number is uniform in all the skeletons which I have examined, with two exceptions, namely, in one of the small feral Porto Santo rabbits and in one of the largest lop-eared kinds; both of these had as usual seven cervical, twelve dorsal with ribs, but, instead of seven lumbar, both had eight lumbar vertebræ. This is remarkable, as Gervais gives seven as the number for the whole genus Lepus. The caudal vertebræ apparently differ by two or three, but I did not attend to them, and they are difficult to count with certainty.