SKULL.

I have carefully examined skulls of ten large lop-eared rabbits, and of five common domestic rabbits, which latter differ from the lop-eared only in not having such large bodies or ears, yet both larger than in the wild rabbit. First for the ten lop-eared rabbits: in all these the skull is remarkably elongated in comparison with its breadth. In a wild rabbit the length was 3.15 inches, in a large fancy rabbit 4.3; whilst the breadth of the cranium enclosing the brain was in both almost exactly the same. Even by taking as the standard of comparison the widest part of the zygomatic arch, the skulls of the lop-eared are proportionally to their breadth three-quarters of an inch too long. The depth of the head has increased almost in the same proportion with the length; it is the breadth alone which has not increased. The parietal and occipital bones enclosing the brain are less arched, both in a longitudinal and transverse line, than in the wild rabbit, so that the shape of the cranium is somewhat different.

The surface is rougher, less cleanly sculptured, and the lines of sutures are more prominent.

Although the skulls of the large lop-eared rabbits in comparison with those of the wild rabbit are much elongated relatively to their breadth, yet, relatively to the size of body, they are far from elongated. The lop-eared rabbits which I examined were, though not fat, more than twice as heavy as the wild specimens; but the skull was very far from being twice as long.

Even if we take the fairer standard of the length of body, from the nose to the a.n.u.s, the skull is not on an average as long as it ought to be by a third of an inch. In the small feral Porto Santo rabbit, on the other hand, the head relatively to the length of body is about a quarter of an inch too long.

This elongation of the skull relatively to its breadth, I find a universal character, not only with the large lop-eared rabbits, but in all the artificial breeds; as is well seen in the skull of the Angora. I was at first much surprised at the fact, and could not imagine why domestication could produce this uniform result; but the explanation seems to lie in the circ.u.mstance that during a number of generations the artificial races have been closely confined, and have had little occasion to exert either their senses, or intellect, or voluntary muscles; consequently the brain, as we shall presently more fully see, has not increased relatively with the size of body. As the brain has not increased, the bony case enclosing it has not increased, and this has evidently affected through correlation the breadth of the entire skull from end to end.

(FIGURE 6. SKULL OF WILD RABBIT, of natural size.

FIGURE 7. SKULL OF LARGE LOP-EARED RABBIT, of natural size.

FIGURE 8. PART OF ZYGOMATIC ARCH, showing the projecting end of the malar bone of the auditory meatus: of natural size. Upper figure, Wild Rabbit.

Lower figure, Lop-eared, hare-coloured Rabbit.

FIGURE 9. POSTERIOR END OF SKULL, of natural size, showing the inter- parietal bone. A. Wild Rabbit. B. Feral Rabbit from island of P. Santo, near Madeira. C. Large Lop-eared Rabbit.)

In all the skulls of the large lop-eared rabbits, the supra-orbital plates or processes of the frontal bones are much broader than in the wild rabbit, and they generally project more upwards. In the zygomatic arch the posterior or projecting point of the malar-bone is broader and blunter; and in the specimen, figure 8, it is so in a remarkable degree. This point approaches nearer to the auditory meatus than in the wild rabbit, as may be best seen in figure 8; but this circ.u.mstance mainly depends on the changed direction of the meatus. The inter-parietal bone (see figure 9) differs much in shape in the several skulls; generally it is more oval, that is more extended in the line of the longitudinal axis of the skull, than in the wild rabbit. The posterior margin of "the square raised platform"

(4/25. Waterhouse 'Nat. Hist. Mammalia' volume 2 page 36.) of the occiput, instead of being truncated, or projecting slightly as in the wild rabbit, is in most lop-eared rabbits pointed, as in figure 9, C. The paramastoids relatively to the size of the skull are generally much thicker than in the wild rabbit.

(FIGURE 10. OCCIPITAL FORAMEN, of natural size, in--A. Wild Rabbit; B.

Large Lop-eared Rabbit.)

The occipital foramen (figure 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 pa.s.sage to so important a structure as the spinal marrow, though apparently the outline of the latter is not affected by the shape of the pa.s.sage.

(FIGURE 11. SKULL, of natural size, of Half-lop Rabbit, showing the different direction of the auditory meatus on the two sides, and the consequent general distortion of the skull. The left ear of the animal (or right side of figure) lopped forwards.)

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 figure 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 (figure 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!

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 ill.u.s.trating the law that varieties of one species often a.s.sume 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.

VERTEBRAE.

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 vertebrae. This is remarkable, as Gervais gives seven as the number for the whole genus Lepus. The caudal vertebrae apparently differ by two or three, but I did not attend to them, and they are difficult to count with certainty.

(FIGURE 12. ATLAS VERTEBRAE, of natural size; inferior surface viewed obliquely. Upper figure, Wild Rabbit. Lower figure, Hare-coloured, large, Lop-eared Rabbit, a, supra-median, atlantoid process; b, infra-median process.)

In the first cervical vertebra, or atlas, the anterior margin of the neural arch varies a little in wild specimens, being either nearly smooth, or furnished with a small supra-median atlantoid process; I have figured a specimen with the largest process (a) which I have seen; but it will be observed how inferior this is in size and different in shape to that in a large lop-eared rabbit. In the latter, the infra-median process (b) is also proportionally much thicker and longer. The alae are a little squarer in outline.

(FIGURE 13. THIRD CERVICAL VERTEBRAE, of natural size, of: A. Wild Rabbit; B. Hare-coloured, large, Lop-eared Rabbit, a, a, inferior surface; b, b, anterior articular surfaces.)

THIRD CERVICAL VERTEBRA.

In the wild rabbit (figure 13, A a) this vertebra, viewed on the inferior surface, has a transverse process, which is directed obliquely backwards, and consists of a single pointed bar; in the fourth vertebra this process is slightly forked in the middle. In the large lop-eared rabbits this process (B a) is forked in the third vertebra, as in the fourth of the wild rabbit. But the third cervical vertebrae of the wild and lop-eared (A b, B b) rabbits differ more conspicuously when their anterior articular surfaces are compared; for the extremities of the antero-dorsal processes in the wild rabbit are simply rounded, whilst in the lop-eared they are trifid, with a deep central pit. The ca.n.a.l for the spinal marrow in the lop-eared (B b) is more elongated in a transverse direction than in the wild rabbit; and the pa.s.sages for the arteries are of a slightly different shape. These several differences in this vertebra seem to me well deserving attention.

FIRST DORSAL VERTEBRA.

Its neural spine varies in length in the wild rabbit; being sometimes very short, but generally more than half as long as that of the second dorsal; but I have seen it in two large lop-eared rabbits three-fourths of the length of that of the second dorsal vertebra.

(FIGURE 14. DORSAL VERTEBRAE, from sixth to tenth inclusive, of natural size, viewed laterally. A. Wild Rabbit. B. Large, Hare-coloured, so called Spanish Rabbit.)

NINTH AND TENTH DORSAL VERTEBRAE.

In the wild rabbit the neural spine of the ninth vertebra is just perceptibly thicker than that of the eighth; and the neural spine of the tenth is plainly thicker and shorter than those of all the anterior vertebrae. In the large lop-eared rabbits the neural spines of the tenth, ninth, and eighth vertebrae, and even in a slight degree that of the seventh, are very much thicker, and of somewhat different shape, in comparison with those of the wild rabbit. So that this part of the vertebral column differs considerably in appearance from the same part in the wild rabbit, and closely resembles in an interesting manner these same vertebrae in some species of hares. In the Angora, Chinchilla, and Himalayan rabbits, the neural spines of the eighth and ninth vertebrae are in a slight degree thicker than in the wild. On the other hand, in one of the feral Porto Santo rabbits, which in most of its characters deviates from the common wild rabbit, in a direction exactly opposite to that a.s.sumed by the large lop-eared rabbits, the neural spines of the ninth and tenth vertebrae were not at all larger than those of the several anterior vertebra. In this same Porto Santo specimen there was no trace in the ninth vertebra of the anterior lateral processes (see figure 14), which are plainly developed in all British wild rabbits, and still more plainly developed in the large lop-eared rabbits. In a half-wild rabbit from Sandon Park (4/26. These rabbits have run wild for a considerable time in Sandon Park, and in other places in Staffordshire and Shropshire. They originated, as I have been informed by the gamekeeper, from variously-coloured domestic rabbits which had been turned out. They vary in colour; but many are symmetrically coloured, being white with a streak along the spine, and with the ears and certain marks about the head of a blackish-grey tint. They have rather longer bodies than common rabbits), a haemal spine was moderately well developed on the under side of the twelfth dorsal vertebra, and I have seen this in no other specimen.

LUMBAR VERTEBRAE.

I have stated that in two cases there were eight instead of seven lumbar vertebrae. The third lumbar vertebrae in one skeleton of a wild British rabbit, and in one of the Porto Santo feral rabbits, had a haemal spine; whilst in four skeletons of large lop-eared rabbits, and in the Himalayan rabbit, this same vertebra had a well developed haemal spine.

PELVIS.

In four wild specimens this bone was almost absolutely identical in shape; but in several domesticated breeds shades of differences could be distinguished. In the large lop-eared rabbits, the whole upper part of the ilium is straighter, or less splayed outwards, than in the wild rabbit; and the tuberosity on the inner lip of the anterior and upper part of the ilium is proportionally more prominent.

(FIGURE 15. TERMINAL BONE OF STERNUM, of natural size, A. Wild Rabbit. B.

Hare-coloured, Lop-eared Rabbit. C. Hare-coloured Spanish Rabbit. (N.B. The left-hand angle of the upper articular extremity of B was broken, and has been accidentally thus represented.))

STERNUM.

The posterior end of the posterior sternal bone in the wild rabbit (figure 15, A) is thin and slightly enlarged; in some of the large lop-eared rabbits (B) it is much more enlarged towards the extremity; whilst in other specimens (C) it keeps nearly of the same breadth from end to end, but is much thicker at the extremity.

(FIGURE 16. ACROMION OF SCAPULA, of natural size. A. Wild Rabbit. B, C, D, Large, Lop-eared Rabbits.)

SCAPULA.

The acromion sends out a rectangular bar, ending in an oblique k.n.o.b, which latter in the wild rabbit (figure 16, A) varies a little in shape and size, as does the apex of the acromion in sharpness, and the part just below the rectangular bar in breadth. But the variations in these respects in the wild rabbit are very slight: whilst in the large lop-eared rabbits they are considerable. Thus in some specimens (B) the oblique terminal k.n.o.b is developed into a short bar, forming an obtuse angle with the rectangular bar. In another specimen (C) these two unequal bars form nearly a straight line. The apex of the acromion varies much in breadth and sharpness, as may be seen by comparing figures B, C, and D.

LIMBS.

In these I could detect no variation; but the bones of the feet were too troublesome to compare with much care.]

I have now described all the differences in the skeletons which I have observed. It is impossible not to be struck with the high degree of variability or plasticity of many of the bones. We see how erroneous the often-repeated statement is, that only the crests of the bones which give attachment to muscles vary in shape, and that only parts of slight importance become modified under domestication. No one will say, for instance, that the occipital foramen, or the atlas, or the third cervical vertebra is a part of slight importance. If the several vertebrae of the wild and lop-eared rabbits, of which figures have been given, had been found fossil, palaeontologists would have declared without hesitation that they had belonged to distinct species.

[THE EFFECTS OF THE USE AND DISUSE OF PARTS.

In the large lop-eared rabbits the relative proportional length of the bones of the same leg, and of the front and hind legs compared with each other, have remained nearly the same as in the wild rabbit; but in weight, the bones of the hind legs apparently have not increased in due proportion with the front legs. The weight of the whole body in the large rabbits examined by me was from twice to twice and a half as great as that of the wild rabbit; and the weight of the bones of the front and hind limbs taken together (excluding the feet, on account of the difficulty of cleaning so many small bones) has increased in the large lop-eared rabbits in nearly the same proportion; consequently in due proportion to the weight of body which they have to support. If we take the length of the body as the standard of comparison, the limbs of the large rabbits have not increased in length in due proportion by one inch and a half. Again, if we take as the standard of comparison the length of the skull, which, as we have before seen, has not increased in length in due proportion to the length of body, the limbs will be found to be, proportionally with those of the wild rabbit, from half to three-quarters of an inch too short. Hence, whatever standard of comparison be taken, the limb-bones of the large lop-eared rabbits have not increased in length, though they have in weight, in full proportion to the other parts of the frame; and this, I presume, may be accounted for by the inactive life which during many generations they have spent. Nor has the scapula increased in length in due proportion to the increased length of the body.

The capacity of the osseous case of the brain is a more interesting point, to which I was led to attend by finding, as previously stated, that with all domesticated rabbits the length of the skull relatively to its breadth has greatly increased in comparison with that of the wild rabbits. If we had possessed a large number of domesticated rabbits of nearly the same size with the wild rabbits, it would have been a simple task to have measured and compared the capacities of their skulls. But this is not the case: almost all the domestic breeds have larger bodies than wild rabbits, and the lop-eared kinds are more than double their weight. As a small animal has to exert its senses, intellect, and instincts equally with a large animal, we ought not by any means to expect an animal twice or thrice as large as another to have a brain of double or treble the size. (4/27.

See Prof. Owen's remarks on this subject in his paper on the 'Zoological Significance of the Brain, etc., of Man, etc.' read before Brit.

a.s.sociation 1862: with respect to Birds see 'Proc. Zoolog. Soc.' January 11, 1848 page 8.) Now, after weighing the bodies of four wild rabbits, and of four large but not fattened lop-eared rabbits, I find that on an average the wild are to the lop-eared in weight as 1 to 2.17; in average length of body as 1 to 1.41; whilst in capacity of skull they are as 1 to 1.15. Hence we see that the capacity of the skull, and consequently the size of the brain, has increased but little, relatively to the increased size of the body; and this fact explains the narrowness of the skull relatively to its length in all domestic rabbits.

TABLE 3: MEASUREMENTS OF WILD AND SEMI-WILD RABBITS.