The Variation of Animals and Plants under Domestication — Volume 1

On the whole we may conclude, more especially from the researches of Boyd Dawkins, that European cattle are descended from two species; and there is no improbability in this fact, for the genus Bos readily yields to domestication. Besides these two species and the zebu, the yak, the gayal, and the arni (3/47. Isid. Geoffroy Saint-Hilaire 'Hist. Nat. Gen.' tome 3, 96.) (not to mention the buffalo or genus Bubalus) have been domesticated; making altogether six species of Bos. The zebu and the two European species are now extinct in a wild state. Although certain races of cattle were domesticated at a very ancient period in Europe, it does not follow that they were first domesticated here. Those who place much reliance on philology argue that they were imported from the East. (3/48. Idem tome 3 pages 82, 91.) It is probable that they originally inhabited a temperate or cold climate, but not a land long covered with snow; for our cattle, as we have seen in the chapter on Horses, have not the instinct of scraping away the snow to get at the herbage beneath. No one could behold the magnificent wild bulls on the bleak Falkland Islands in the southern hemisphere, and doubt about the climate being admirably suited to them. Azara has remarked that in the temperate regions of La Plata the cows conceive when two years old, whilst in the much hotter country of Paraguay they do not conceive till three years old; "from which fact," as he adds, "one may conclude that cattle do not succeed so well in warm countries." (3/49. 'Quadrupedes du Paraguay' tome 2 page 360.)

Bos primigenius and longifrons have been ranked by nearly all palaeontologists as distinct species; and it would not be reasonable to take a different view simply because their domesticated descendants now intercross with the utmost freedom. All the European breeds have so often been crossed both intentionally and unintentionally, that, if any sterility had ensued from such unions, it would certainly have been detected. As zebus inhabit a distant and much hotter region, and as they differ in so many characters from our European cattle, I have taken pains to ascertain whether the two forms are fertile when crossed. The late Lord Powis imported some zebus and crossed them with common cattle in Shropshire; and I was assured by his steward that the cross-bred animals were perfectly fertile with both parent-stocks. Mr. Blyth informs me that in India hybrids, with various proportions of either blood, are quite fertile; and this can hardly fail to be known, for in some districts (3/50. Walther 'Das Rindvieh' 1817 s. 30.) the two species are allowed to breed freely together. Most of the cattle which were first introduced into Tasmania were humped, so that at one time thousands of crossed animals existed there; and Mr. B. O'Neile Wilson, M.A., writes to me from Tasmania that he has never heard of any sterility having been observed. He himself formerly possessed a herd of such crossed cattle, and all were perfectly fertile; so much so, that he cannot remember even a single cow failing to calve. These several facts afford an important confirmation of the Pallasian doctrine that the descendants of species which when first domesticated would if crossed have been in all probability in some degree sterile, become perfectly fertile after a long course of domestication. In a future chapter we shall see that this doctrine throws some light on the difficult subject of Hybridism.

I have alluded to the cattle in Chillingham Park, which, according to Rutimeyer, have been very little changed from the Bos primigenius type. This park is so ancient that it is referred to in a record of the year 1220. The cattle in their instincts and habits are truly wild. They are white, with the inside of the ears reddish-brown, eyes rimmed with black, muzzles brown, hoofs black, and horns white tipped with black. Within a period of thirty-three years about a dozen calves were born with "brown and blue spots upon the cheeks or necks; but these, together with any defective animals, were always destroyed." According to Bewick, about the year 1770 some calves appeared with black ears; but these were also destroyed by the keeper, and black ears have not since reappeared. The wild white cattle in the Duke of Hamilton's park, where I have heard of the birth of a black calf, are said by Lord Tankerville to be inferior to those at Chillingham. The cattle kept until the year 1780 by the Duke of Queensberry, but now extinct, had their ears, muzzle, and orbits of the eyes black. Those which have existed from time immemorial at Chartley, closely resemble the cattle at Chillingham, but are larger, "with some small difference in the colour of the ears." "They frequently tend to become entirely black; and a singular superstition prevails in the vicinity that, when a black calf is born, some calamity impends over the noble house of Ferrers. All the black calves are destroyed." The cattle at Burton Constable in Yorkshire, now extinct, had ears, muzzle, and the tip of the tail black. Those at Gisburne, also in Yorkshire, are said by Bewick to have been sometimes without dark muzzles, with the inside alone of the ears brown; and they are elsewhere said to have been low in stature and hornless. (3/51. I am much indebted to the present Earl of Tankerville for information about his wild cattle; and for the skull which was sent to Prof. Rutimeyer. The fullest account of the Chillingham cattle is given by Mr. Hindmarsh, together with a letter by the late Lord Tankerville, in 'Annals and Mag. of Nat. Hist.' volume 2 1839 page 274. See Bewick 'Quadrupeds' 2nd edition 1791 page 35 note. With respect to those of the Duke of Queensberry see Pennant 'Tour in Scotland' page 109. For those of Chartley, see Low 'Domesticated Animals of Britain' 1845 page 238. For those of Gisburne see Bewick 'Quadrupeds' and 'Encyclop. of Rural Sports' page 101.)

The several above-specified differences in the park-cattle, slight though they be, are worth recording, as they show that animals living nearly in a state of nature, and exposed to nearly uniform conditions, if not allowed to roam freely and to cross with other herds, do not keep as uniform as truly wild animals. For the preservation of a uniform character, even within the same park, a certain degree of selection — that is, the destruction of the dark-coloured calves — is apparently necessary.

Boyd Dawkins believes that the park-cattle are descended from anciently domesticated, and not truly wild animals; and from the occasional appearance of dark-coloured calves, it is improbable that the aboriginal Bos primigenius was white. It is curious what a strong, though not invariable, tendency there is in wild or escaped cattle to become white with coloured ears, under widely different conditions of life. If the old writers Boethius and Leslie (3/52. Boethius was born in 1470; 'Annals and Mag. of Nat. Hist.' volume 2 1839 page 281; and volume 4 1849 page 424.) can be trusted, the wild cattle of Scotland were white and furnished with a great mane; but the colour of their ears is not mentioned. In Wales (3/53. 'Youatt on Cattle' 1834 page 48: See also page 242, on shorthorn cattle. Bell in his 'British Quadrupeds' page 423 states that, after long attending to the subject, he has found that white cattle invariably have coloured ears.), during the tenth century, some of the cattle are described as being white with red ears. Four hundred cattle thus coloured were sent to King John; and an early record speaks of a hundred cattle with red ears having been demanded as a compensation for some offence, but, if the cattle were of a dark or black colour, 150 were to be presented. The black cattle of North Wales apparently belong, as we have seen, to the small longifrons type: and as the alternative was offered of either 150 dark cattle, or 100 white cattle with red ears, we may presume that the latter were the larger beasts, and probably belonged to the primigenius type. Youatt has remarked that at the present day, whenever cattle of the shorthorn breed are white, the extremities of their ears are more or less tinged with red.

The cattle which have run wild on the Pampas, in Texas, and in two parts of Africa, have become of a nearly uniform dark brownish-red. (3/54. Azara 'Quadrupedes du Paraguay' tome 2 page 361. Azara quotes Buffon for the feral cattle of Africa. For Texas see 'Times' February 18, 1846.) On the Ladrone Islands, in the Pacific Ocean, immense herds of cattle, which were wild in the year 1741, are described as "milk-white, except their ears, which are generally black." (3/55. Anson's Voyage. See Kerr and Porter 'Collection' volume 12 page 103.) The Falkland Islands, situated far south, with all the conditions of life as different as it is possible to conceive from those of the Ladrones, offer a more interesting case. Cattle have run wild there during eighty or ninety years; and in the southern districts the animals are mostly white, with their feet, or whole heads, or only their ears black; but my informant, Admiral Sulivan (3/56. See also Mr. Mackinnon's pamphlet on the Falkland Islands, page 24.), who long resided on these islands, does not believe that they are ever purely white. So that in these two archipelagos we see that the cattle tend to become white with coloured ears. In other parts of the Falkland Islands other colours prevail: near Port Pleasant brown is the common tint; round Mount Usborn, about half the animals in some of the herds were lead- or mouse-coloured, which elsewhere is an unusual tint. These latter cattle, though generally inhabiting high land, breed about a month earlier than the other cattle; and this circumstance would aid in keeping them distinct and in perpetuating a peculiar colour. It is worth recalling to mind that blue or lead-coloured marks have occasionally appeared on the white cattle of Chillingham. So plainly different were the colours of the wild herds in different parts of the Falkland Islands, that in hunting them, as Admiral Sulivan informs me, white spots in one district, and dark spots in another district, were always looked out for on the distant hills. In the intermediate districts, intermediate colours prevailed. Whatever the cause may be, this tendency in the wild cattle of the Falkland Islands, which are all descended from a few brought from La Plata, to break up into herds of three different colours, is an interesting fact.

Returning to the several British breeds, the conspicuous difference in general appearance between Shorthorns, Longhorns (now rarely seen), Herefords, Highland cattle, Alderneys, etc., must be familiar to every one. A part of this difference may be attributed to descent from primordially distinct species; but we may feel sure that there has been a considerable amount of variation. Even during the Neolithic period, the domestic cattle were to a certain extent variable. Within recent times most of the breeds have been modified by careful and methodical selection. How strongly the characters thus acquired are inherited, may be inferred from the prices realised by the improved breeds; even at the first sale of Colling's Shorthorns, eleven bulls reached an average of 214 pounds, and lately Shorthorn bulls have been sold for a thousand guineas, and have been exported to all quarters of the world.

Some constitutional differences may be here noticed. The Shorthorns arrive at maturity far earlier than the wilder breeds, such as those of Wales or the Highlands. This fact has been shown in an interesting manner by Mr. Simonds (3/57. 'The Age of the Ox, Sheep, Pig' etc. by Prof. James Simonds, published by order of the Royal Agricult. Soc.) who has given a table of the average period of their dentition, which proves that there is a difference of no less than six months in the appearance of the permanent incisors. The period of gestation, from observations made by Tessier on 1131 cows, varies to the extent of eighty-one days; and what is more interesting, M. Lefour affirms "that the period of gestation is longer in the large German cattle than in the smaller breeds." (3/58. 'Ann. Agricult. France' April 1837 as quoted in 'The Veterinary' volume 12 page 725. I quote Tessier's observations from 'Youatt on Cattle' page 527.) With respect to the period of conception, it seems certain that Alderney and Zetland cows often become pregnant earlier than other breeds. (3/59. 'The Veterinary' volume 8 page 681 and volume 10 page 268. Low 'Domest. Animals, etc.' page 297.) Lastly, as four fully developed mammae is a generic character in the genus Bos (3/60. Mr. Ogleby in 'Proc. Zoolog. Soc.' 1836 page 138, and 1840 page 4. Quatrefages quotes Philippi 'Revue des Cours Scientifiques' February 12, 1688 page 657, that the cattle of Piacentino have thirteen dorsal vertebrae and ribs in the place of the ordinary number of twelve.), it is worth notice that with our domestic cows the two rudimentary mammae often become fairly well developed and yield milk.

As numerous breeds are generally found only in long-civilised countries, it may be well to show that in some countries inhabited by barbarous races, who are frequently at war with each other, and therefore have little free communication, several distinct breeds of cattle now exist or formerly existed. At the Cape of Good Hope Leguat observed, in the year 1720, three kinds. (3/61. Leguat's Voyage quoted by Vasey in his 'Delineations of the Ox-tribe' page 132.) At the present day various travellers have noticed the differences in the breeds in Southern Africa. Sir Andrew Smith several years ago remarked to me that the cattle possessed by the different tribes of Caffres, though living near each other under the same latitude and in the same kind of country, yet differed, and he expressed much surprise at the fact. Mr. Andersson has described (3/62. 'Travels in South Africa' pages 317, 336.) the Damara, Bechuana, and Namaqua cattle; and he informs me in a letter that the cattle north of Lake Ngami are likewise different, as Mr. Galton has heard is also the case with the cattle of Benguela. The Namaqua cattle in size and shape nearly resemble European cattle, and have short stout horns and large hoofs. The Damara cattle are very peculiar, being big-boned, with slender legs, and small hard feet; their tails are adorned with a tuft of long bushy hair nearly touching the ground, and their horns are extraordinarily large. The Bechuana cattle have even larger horns, and there is now a skull in London with the two horns 8 ft. 8 1/4 in. long, as measured in a straight line from tip to tip, and no less than 13 ft. 5 in. as measured along their curvature! Mr. Andersson in his letter to me says that, though he will not venture to describe the differences between the breeds belonging to the many different sub-tribes, yet such certainly exist, as shown by the wonderful facility with which the natives discriminate them.

That many breeds of cattle have originated through variation, independently of descent from distinct species, we may infer from what we see in South America, where the genus Bos was not endemic, and where the cattle which now exist in such vast numbers are the descendants of a few imported from Spain and Portugal. In Columbia, Roulin (3/63. 'Mem. de 1'Institut present. par divers Savans' tome 6 1835 page 333. For Brazil see 'Comptes Rendus' June 15, 1846. See Azara 'Quadrupedes du Paraguay' tome 2 pages 359, 361.) describes two peculiar breeds, namely, pelones, with extremely thin and fine hair, and calongos, absolutely naked. According to Castelnau there are two races in Brazil, one like European cattle, the other different, with remarkable horns. In Paraguay, Azara describes a breed which certainly originated in S. America, called chivos, "because they have straight vertical horns, conical, and very large at the base." He likewise describes a dwarf race in Corrientes, with short legs and a body larger than usual. Cattle without horns, and others with reversed hair, have also originated in Paraguay.

Another monstrous breed, called niatas or natas, of which I saw two small herds on the northern bank of the Plata, is so remarkable as to deserve a fuller description. This breed bears the same relation to other breeds, as bull or pug dogs do to other dogs, or as improved pigs, according to H. von Nathusius, do to common pigs. (3/64. 'Schweineschadel' 1864 s. 104. Nathusius states that the form of skull characteristic in the niata cattle occasionally appears in European cattle; but he is mistaken, as we shall hereafter see, in supposing that these cattle do not form a distinct race. Prof. Wyman, of Cambridge, United States, informs me that the common cod- fish presents a similar monstrosity, called by the fishermen "bull-dog cod." Prof. Wyman also concluded, after making numerous inquiries in La Plata, that the niata cattle transmit their peculiarities or form a race.) Rutimeyer believes that these cattle belong to the primigenius type. (3/65 'Ueber Art des zahmen Europ. Rindes' 1866 s. 28.) The forehead is very short and broad, with the nasal end of the skull, together with the whole plane of the upper molar-teeth, curved upwards. The lower jaw projects beyond the upper, and has a corresponding upward curvature. It is an interesting fact that an almost similar confirmation characterizes, as I am informed by Dr. Falconer, the extinct and gigantic Sivatherium of India, and is not known in any other ruminant. The upper lip is much drawn back, the nostrils are seated high up and are widely open, the eyes project outwards, and the horns are large. In walking the head is carried low, and the neck is short. The hind legs appear to be longer, compared with the front legs, than is usual. The exposed incisor teeth, the short head and upturned nostrils, give these cattle the most ludicrous, self-confident air of defiance. The skull which I presented to the College of Surgeons has been thus described by Professor Owen (3/66. 'Descriptive Cat. of Ost. Collect. of College of Surgeons' 1853 page 624. Vasey in his 'Delineations of the Ox-tribe' has given a figure of this skull; and I sent a photograph of it to Prof. Rutimeyer.) "It is remarkable from the stunted development of the nasals, premaxillaries, and fore-part of the lower jaw, which is unusually curved upwards to come into contact with the premaxillaries. The nasal bones are about one-third the ordinary length, but retain almost their normal breadth. The triangular vacuity is left between them, the frontal and lachrymal, which latter bone articulates with the premaxillary, and thus excludes the maxillary from any junction with the nasal." So that even the connexion of some of the bones is changed. Other differences might be added: thus the plane of the condyles is somewhat modified, and the terminal edge of the premaxillaries forms an arch. In fact, on comparison with the skull of a common ox, scarcely a single bone presents the same exact shape, and the whole skull has a wonderfully different appearance.

The first brief published notice of this race was by Azara, between the years 1783-96; but Don F. Muniz, of Luxan, who has kindly collected information for me, states that about 1760 these cattle were kept as curiosities near Buenos Ayres. Their origin is not positively known, but they must have originated subsequently to the year 1552, when cattle were first introduced. Senor Muniz informs me that the breed is believed to have originated with the Indians southward of the Plata. Even to this day those reared near the Plata show their less civilised nature in being fiercer than common cattle, and in the cow, if visited too often, easily deserting her first calf. The breed is very true, and a niata bull and cow invariably produce niata calves. The breed has already lasted at least a century. A niata bull crossed with a common cow, and the reverse cross, yield offspring having an intermediate character, but with the niata character strongly displayed. According to Senor Muniz, there is the clearest evidence, contrary to the common belief of agriculturists in analogous cases, that the niata cow when crossed with a common bull transmits her peculiarities more strongly than does the niata bull when crossed with a common cow. When the pasture is tolerably long, these cattle feed as well as common cattle with their tongue and palate; but during the great droughts, when so many animals perish on the Pampas, the niata breed lies under a great disadvantage, and would, if not attended to, become extinct; for the common cattle, like horses, are able to keep alive by browsing with their lips on the twigs of trees and on reeds: this the niatas cannot so well do, as their lips do not join, and hence they are found to perish before the common cattle. This strikes me as a good illustration of how little we are able to judge from the ordinary habits of an animal, on what circumstances, occurring only at long intervals of time, its rarity or extinction may depend. It shows us, also, how natural selection would have determined the rejection of the niata modification had it arisen in a state of nature.

Having described the semi-monstrous niata breed, I may allude to a white bull, said to have been brought from Africa, which was exhibited in London in 1829, and which has been well figured by Mr. Harvey. (3/67. Loudon's 'Magazine of Nat. Hist.' volume 1 1829 page 113. Separate figures are given of the animal, its hoofs, eye, and dewlap.) It had a hump, and was furnished with a mane. The dewlap was peculiar, being divided between its fore-legs into parallel divisions. Its lateral hoofs were annually shed, and grew to the length of five or six inches. The eye was very peculiar, being remarkably prominent, and "resembled a cup and ball, thus enabling the animal to see on all sides with equal ease; the pupil was small and oval, or rather a parallelogram with the ends cut off, and lying transversely across the ball." A new and strange breed might probably have been formed by careful breeding and selection from this animal.

I have often speculated on the probable causes through which each separate district in Great Britain came to possess in former times its own peculiar breed of cattle; and the question is, perhaps, even more perplexing in the case of Southern Africa. We now know that the differences may be in part attributed to descent from distinct species; but this cause is far from sufficient. Have the slight differences in climate and in the nature of the pasture, in the different districts of Britain, directly induced corresponding differences in the cattle? We have seen that the semi-wild cattle in the several British parks are not identical in colouring or size, and that some degree of selection has been requisite to keep them true. It is almost certain that abundant food given during many generations directly affects the size of a breed. (3/68. Low 'Domesticated Animals of the British Isles' page 264.) That climate directly affects the thickness of the skin and the hair is likewise certain: thus Roulin asserts (69 'Mem. de l'Institut present. Par divers Savans' tome 6 1835 page 332.) that the hides of the feral cattle on the hot Llanos "are always much less heavy than those of the cattle raised on the high platform of Bogota; and that these hides yield in weight and in thickness of hair to those of the cattle which have run wild on the lofty Paramos." The same difference has been observed in the hides of the cattle reared on the bleak Falkland Islands and on the temperate Pampas. Low has remarked (3/70. Idem pages 304, 368 etc.) that the cattle which inhabit the more humid parts of Britain have longer hair and thicker skins than other British cattle. When we compare highly improved stall-fed cattle with the wilder breeds, or compare mountain and lowland breeds, we cannot doubt that an active life, leading to the free use of the limbs and lungs, affects the shape and proportions of the whole body. It is probable that some breeds, such as the semi- monstrous niata cattle, and some peculiarities, such as being hornless, etc., have appeared suddenly owing to what we may call in our ignorance spontaneous variation; but even in this case a rude kind of selection is necessary, and the animals thus characterised must be at least partially separated from others. This degree of care, however, has sometimes been taken even in little-civilised districts, where we should least have expected it, as in the case of the niata, chivo, and hornless cattle in S. America.

That methodical selection has done wonders within a recent period in modifying our cattle, no one doubts. During the process of methodical selection it has occasionally happened that deviations of structure, more strongly pronounced than mere individual differences, yet by no means deserving to be called monstrosities, have been taken advantage of: thus the famous Longhorn Bull, Shakespeare, though of the pure Canley stock, "scarcely inherited a single point of the long-horned breed, his horns excepted (3/71. 'Youatt on Cattle' page 193. A full account of this bull is taken from Marshall.); yet in the hands of Mr. Fowler, this bull greatly improved his race. We have also reason to believe that selection, carried on so far unconsciously that there was at no one time any distinct intention to improve or change the breed, has in the course of time modified most of our cattle; for by this process, aided by more abundant food, all the lowland British breeds have increased greatly in size and in early maturity since the reign of Henry VII. (3/72. 'Youatt on Cattle' page 116. Lord Spencer has written on this same subject.) It should never be forgotten that many animals have to be annually slaughtered; so that each owner must determine which shall be killed and which preserved for breeding. In every district, as Youatt has remarked, there is a prejudice in favour of the native breed; so that animals possessing qualities, whatever they may be, which are most valued in each district, will be oftenest preserved; and this unmethodical selection assuredly will in the long run affect the character of the whole breed. But it may be asked, can this rude kind of selection have been practised by barbarians such as those of southern Africa? In a future chapter on Selection we shall see that this has certainly occurred to some extent. Therefore, looking to the origin of the many breeds of cattle which formerly inhabited the several districts of Britain, I conclude that, although slight differences in the nature of the climate, food, etc., as well as changed habits of life, aided by correlation of growth, and the occasional appearance from unknown causes of considerable deviations of structure, have all probably played their parts; yet that the occasional preservation in each district of those individual animals which were most valued by each owner has perhaps been even more effective in the production of the several British breeds. As soon as two or more breeds were formed in any district, or when new breeds descended from distinct species were introduced, their crossing, especially if aided by some selection, will have multiplied the number and modified the characters of the older breeds.

SHEEP.

I shall treat this subject briefly. Most authors look at our domestic sheep as descended from several distinct species. Mr. Blyth, who has carefully attended to the subject, believes that fourteen wild species now exist, but "that not one of them can be identified as the progenitor of any one of the interminable domestic races." M. Gervais thinks that there are six species of Ovis (3/73. Blyth on the genus Ovis in 'Annals and Mag. of Nat. History' volume 7 1841 page 261. With respect to the parentage of the breeds see Mr. Blyth's excellent articles in 'Land and Water' 1867 pages 134, 156. Gervais 'Hist. Nat. des Mammiferes' 1855 tome 2 page 191.) but that our domestic sheep form a distinct genus, now completely extinct. A German naturalist (3/74. Dr. L. Fitzinger 'Ueber die Racen des Zahmen Schafes' 1860 s. 86.) believes that our sheep descend from ten aboriginally distinct species, of which only one is still living in a wild state! Another ingenious observer (3/75. J. Anderson 'Recreations in Agriculture and Natural History' volume 2 page 264.), though not a naturalist, with a bold defiance of everything known on geographical distribution, infers that the sheep of Great Britain alone are the descendants of eleven endemic British forms! Under such a hopeless state of doubt it would be useless for my purpose to give a detailed account of the several breeds; but a few remarks may be added.

Sheep have been domesticated from a very ancient period. Rutimeyer (3/76. 'Pfahlbauten' s. 127, 193.) found in the Swiss lake-dwellings the remains of a small breed, with thin tall legs, and horns like those of a goat, thus differing somewhat from any kind now known. Almost every country has its own peculiar breed; and many countries have several breeds differing greatly from each other. One of the most strongly marked races is an Eastern one with a long tail, including, according to Pallas, twenty vertebrae, and so loaded with fat that it is sometimes placed on a truck, which is dragged about by the living animal. These sheep, though ranked by Fitzinger as a distinct aboriginal form, bear in their drooping ears the stamp of long domestication. This is likewise the case with those sheep which have two great masses of fat on the rump, with the tail in a rudimentary condition. The Angola variety of the long-tailed race has curious masses of fat on the back of the head and beneath the jaws. (3/77. 'Youatt on Sheep' page 120.) Mr. Hodgson in an admirable paper (3/78. 'Journal of the Asiatic Soc. of Bengal' volume 16 pages 1007, 1016.) on the sheep of the Himalaya infers from the distribution of the several races, "that this caudal augmentation in most of its phases is an instance of degeneracy in these pre-eminently Alpine animals." The horns present an endless diversity in character; being not rarely absent, especially in the female sex, or, on the other hand, amounting to four or even eight in number. The horns, when numerous, arise from a crest on the frontal bone, which is elevated in a peculiar manner. It is remarkable that multiplicity of horns "is generally accompanied by great length and coarseness of the fleece." (3/79. 'Youatt on Sheep' pages 142-169.) This correlation, however, is far from being general; for instance, I am informed by Mr. D. Forbes, that the Spanish sheep in Chile resemble, in fleece and in all other characters, their parent merino-race, except that instead of a pair they generally bear four horns. The existence of a pair of mammae is a generic character in the genus Ovis as well as in several allied forms; nevertheless, as Mr. Hodgson has remarked, "this character is not absolutely constant even among the true and proper sheep: for I have more than once met with Cagias (a sub-Himalayan domestic race) possessed of four teats." (3/80. 'Journal Asiat. Soc. of Bengal' volume 16 1847 page 1015.) This case is the more remarkable as, when any part or organ is present in reduced number in comparison with the same part in allied groups, it usually is subject to little variation. The presence of interdigital pits has likewise been considered as a generic distinction in sheep; but Isidore Geoffroy (3/81. 'Hist. Nat. Gen.' tome 3 page 435.) has shown that these pits or pouches are absent in some breeds.

In sheep there is a strong tendency for characters, which have apparently been acquired under domestication, to become attached either exclusively to the male sex, or to be more highly developed in this than in the other sex. Thus in many breeds the horns are deficient in the ewe, though this likewise occurs occasionally with the female of the wild musmon. In the rams of the Wallachian breed, "the horns spring almost perpendicularly from the frontal bone, and then take a beautiful spiral form; in the ewes they protrude nearly at right angles from the head, and then become twisted in a singular manner." (3/82. 'Youatt on Sheep' page 138.) Mr. Hodgson states that the extraordinarily arched nose or chaffron, which is so highly developed in several foreign breeds, is characteristic of the ram alone, and apparently is the result of domestication. (3/83. 'Journal Asiat. Soc. of Bengal' volume 16 1847 pages 1015, 1016.) I hear from Mr. Blyth that the accumulation of fat in the fat-tailed sheep of the plains of India is greater in the male than in the female; and Fitzinger (3/84. 'Racen des Zahmen Schafes' s. 77.) remarks that the mane in the African maned race is far more developed in the ram than in the ewe.

Different races of sheep, like cattle, present constitutional differences. Thus the improved breeds arrive at maturity at an early age, as has been well shown by Mr. Simonds through their early average period of dentition. The several races have become adapted to different kinds of pasture and climate: for instance, no one can rear Leicester sheep on mountainous regions, where Cheviots flourish. As Youatt has remarked, "In all the different districts of Great Britain we find various breeds of sheep beautifully adapted to the locality which they occupy. No one knows their origin; they are indigenous to the soil, climate, pasturage, and the locality on which they graze; they seem to have been formed for it and by it." (3/85. 'Rural Economy of Norfolk' volume 2 page 136.) Marshall relates (3/86. 'Youatt on Sheep' page 312. On same subject, see excellent remarks in 'Gardener's Chronicle' 1858 page 868. For experiments in crossing Cheviot sheep with Leicesters see Youatt page 325.) that a flock of heavy Lincolnshire and light Norfolk sheep which had been bred together in a large sheep-walk, part of which was low, rich, and moist, and another part high and dry, with benty grass, when turned out, regularly separated from each other; the heavy sheep drawing off to the rich soil, and the lighter sheep to their own soil; so that "whilst there was plenty of grass the two breeds kept themselves as distinct as rooks and pigeons." Numerous sheep from various parts of the world have been brought during a long course of years to the Zoological Gardens of London; but as Youatt, who attended the animals as a veterinary surgeon, remarks, "few or none die of the rot, but they are phthisical; not one of them from a torrid climate lasts out the second year, and when they die their lungs are tuberculated." (3/87. 'Youatt on Sheep' note page 491.) There is very good evidence that English breeds of sheep will not succeed in France. (3/88. M. Malingie-Nouel 'Journal R. Agricult. Soc.' volume 14 1853 page 214 translated and therefore approved by a great authority, Mr. Pusey.) Even in certain parts of England it has been found impossible to keep certain breeds of sheep; thus on a farm on the banks of the Ouse, the Leicester sheep were so rapidly destroyed by pleuritis (3/89. 'The Veterinary' volume 10 page 217.) that the owner could not keep them; the coarser-skinned sheep never being affected.

The period of gestation was formerly thought to be of so unalterable a character, that a supposed difference of this kind between the wolf and the dog was esteemed a sure sign of specific distinction; but we have seen that the period is shorter in the improved breeds of the pig, and in the larger breeds of the ox, than in other breeds of these two animals. And now we know, on the excellent authority of Hermann von Nathusius (3/90. A translation of his paper is given in 'Bull. Soc. Imp. d'Acclimat.' tome 9 1862 page 723.), that Merino and Southdown sheep, when both have long been kept under exactly the same conditions, differ in their average period of gestation, as is seen in the following Table Merinos 150.3 days. Southdowns 144.2 " Half-bred Merinos and Southdowns 146.3 " 3/4 blood of Southdown 145.5 " 7/8 blood of Southdown 144.2 "

In this graduated difference in cross-bred animals having different proportions of Southdown blood, we see how strictly the two periods of gestation have been transmitted. Nathusius remarks that, as Southdowns grow with remarkable rapidity after birth, it is not surprising that their foetal development should have been shortened. It is of course possible that the difference in these two breeds may be due to their descent from distinct parent-species; but as the early maturity of the Southdowns has long been carefully attended to by breeders, the difference is more probably the result of such attention. Lastly, the fecundity of the several breeds differs much; some generally producing twins or even triplets at a birth, of which fact the curious Shangai sheep (with their truncated and rudimentary ears, and great Roman noses), lately exhibited in the Zoological Gardens, offer a remarkable instance.

Sheep are perhaps more readily affected by the direct action of the conditions of life to which they have been exposed than almost any other domestic animal. According to Pallas, and more recently according to Erman, the fat-tailed Kirghisian sheep, when bred for a few generations in Russia, degenerate, and the mass of fat dwindles away, "the scanty and bitter herbage of the steppes seems so essential to their development." Pallas makes an analogous statement with respect to one of the Crimean breeds. Burnes states that the Karakool breed, which produces a fine, curled, black, and valuable fleece, when removed from its own canton near Bokhara to Persia or to other quarters, loses its peculiar fleece. (3/91. Erman 'Travels in Siberia' English translation volume 1 page 228. For Pallas on the fat-tailed sheep I quote from Anderson's account of the 'Sheep of Russia' 1794 page 34. With respect to the Crimean sheep see Pallas 'Travels' English translation volume 2 page 454. For the Karakool sheep see Burnes 'Travels in Bokhara' volume 3 page 151.) In all such cases, however, it may be that a change of any kind in the conditions of life causes variability and consequent loss of character, and not that certain conditions are necessary for the development of certain characters.

Great heat, however, seems to act directly on the fleece: several accounts have been published of the change which sheep imported from Europe undergo in the West Indies. Dr. Nicholson of Antigua informs me that, after the third generation, the wool disappears from the whole body, except over the loins; and the animal then appears like a goat with a dirty door-mat on its back. A similar change is said to take place on the west coast of Africa. (3/92. See Report of the Directors of the Sierra Leone Company as quoted in White 'Gradation of Man' page 95. With respect to the change which sheep undergo in the West Indies see also Dr. Davy in 'Edin. New. Phil. Journal' January 1852. For the statement made by Roulin see 'Mem. de l'Institut present. par divers Savans.' tome 6 1835 page 347.) On the other hand, many wool-bearing sheep live on the hot plains of India. Roulin asserts that in the lower and heated valleys of the Cordillera, if the lambs are sheared as soon as the wool has grown to a certain thickness, all goes on afterwards as usual; but if not sheared, the wool detaches itself in flakes, and short shining hair like that on a goat is produced ever afterwards. This curious result seems merely to be an exaggerated tendency natural to the Merino breed, for as a great authority, namely, Lord Somerville, remarks, "the wool of our Merino sheep after shear-time is hard and coarse to such a degree as to render it almost impossible to suppose that the same animal could bear wool so opposite in quality, compared to that which has been clipped from it: as the cold weather advances, the fleeces recover their soft quality." As in sheep of all breeds the fleece naturally consists of longer and coarser hair covering shorter and softer wool, the change which it often undergoes in hot climates is probably merely a case of unequal development; for even with those sheep which like goats are covered with hair, a small quantity of underlying wool may always be found. (3/93. 'Youatt on Sheep' page 69 where Lord Somerville is quoted. See page 117 on the presence of wool under the hair. With respect to the fleeces of Australian sheep page 185. On selection counteracting any tendency to change see pages 70, 117, 120, 168.) In the wild mountain-sheep (0vis montana) of North America there is an analogous annual change of coat; "the wool begins to drop out in early spring, leaving in its place a coat of hair resembling that of the elk, a change of pelage quite different in character from the ordinary thickening of the coat or hair, common to all furred animals in winter, — for instance, in the horse, the cow, etc., which shed their winter coat in the spring." (3/94. Audubon and Bachman 'The Quadrupeds of North America' 1846 volume 5 page 365.)

A slight difference in climate or pasture sometimes slightly affects the fleece, as has been observed even in different districts in England, and is well shown by the great softness of the wool brought from Southern Australia. But it should be observed, as Youatt repeatedly insists, that the tendency to change may generally be counteracted by careful selection. M. Lasterye, after discussing this subject, sums up as follows: "The preservation of the Merino race in its utmost purity at the Cape of Good Hope, in the marshes of Holland, and under the rigorous climate of Sweden, furnishes an additional support of this my unalterable principle, that fine-woolled sheep may be kept wherever industrious men and intelligent breeders exist."

That methodical selection has effected great changes in several breeds of sheep no one who knows anything on the subject, entertains a doubt. The case of the Southdowns, as improved by Ellman, offers perhaps the most striking instance. Unconscious or occasional selection has likewise slowly produced a great effect, as we shall see in the chapters on Selection. That crossing has largely modified some breeds, no one who will study what has been written on this subject — for instance, Mr. Spooner's paper — will dispute; but to produce uniformity in a crossed breed, careful selection and "rigorous weeding," as this author expresses it, are indispensable. (3/95. 'Journal of R. Agricult. Soc. of England' volume 20 part 2, W.C. Spooner on cross-Breeding.)

In some few instances new breeds have suddenly originated; thus, in 1791, a ram-lamb was born in Massachusetts, having short crooked legs and a long back, like a turnspit-dog. From this one lamb the otter or ancon semi- monstrous breed was raised; as these sheep could not leap over the fences, it was thought that they would be valuable; but they have been supplanted by merinos, and thus exterminated. The sheep are remarkable from transmitting their character so truly that Colonel Humphreys (3/96. 'Philosoph. Transactions' London 1813 page 88.) never heard of "but one questionable case" of an ancon ram and ewe not producing ancon offspring. When they are crossed with other breeds the offspring, with rare exceptions, instead of being intermediate in character, perfectly resemble either parent; even one of twins has resembled one parent and the second the other. Lastly, "the ancons have been observed to keep together, separating themselves from the rest of the flock when put into enclosures with other sheep."

A more interesting case has been recorded in the Report of the Juries for the Great Exhibition (1851), namely, the production of a merino ram-lamb on the Mauchamp farm, in 1828, which was remarkable for its long, smooth, straight, and silky wool. By the year 1833 M. Graux had raised rams enough to serve his whole flock, and after a few more years he was able to sell stock of his new breed. So peculiar and valuable is the wool, that it sells at 25 per cent above the best merino wool: even the fleeces of half-bred animals are valuable, and are known in France as the "Mauchamp-merino." It is interesting, as showing how generally any marked deviation of structure is accompanied by other deviations, that the first ram and his immediate offspring were of small size, with large heads, long necks, narrow chests, and long flanks; but these blemishes were removed by judicious crosses and selection. The long smooth wool was also correlated with smooth horns; and as horns and hair are homologous structures, we can understand the meaning of this correlation. If the Mauchamp and ancon breeds had originated a century or two ago, we should have had no record of their birth; and many a naturalist would no doubt have insisted, especially in the case of the Mauchamp race, that they had each descended from, or been crossed with, some unknown aboriginal form.

GOATS.

From the recent researches of M. Brandt, most naturalists now believe that all our goats are descended from the Capra aegagrus of the mountains of Asia, possibly mingled with the allied Indian species C. falconeri of India. (3/97. Isidore Geoffroy St. Hilaire 'Hist. Nat. Generale' tome 3 page 87. Mr. Blyth 'Land and Water' 1867 page 37 has arrived at a similar conclusion, but he thinks that certain Eastern races may perhaps be in part descended from the Asiatic markhor.) In Switzerland, during the neolithic period, the domestic goat was commoner than the sheep; and this very ancient race differed in no respect from that now common in Switzerland. (3/98. Rutimeyer 'Pfahlbauten' s. 127.) At the present time, the many races found in several parts of the world differ greatly from each other; nevertheless, as far as they have been tried (3/99. Godron 'De l'Espece' tome 1 page 402.) they are all quite fertile when crossed. So numerous are the breeds, that Mr. G. Clark (3/100. 'Annals and Mag. of Nat History' 2nd series volume 2 1848 page 363.) has described eight distinct kinds imported into the one island of Mauritius. The ears of one kind were enormously developed, being, as measured by Mr. Clark, no less than 19 inches in length and 4 3/4 inches in breadth. As with cattle, the mammae of those breeds which are regularly milked become greatly developed; and, as Mr. Clark remarks, "it is not rare to see their teats touching the ground." The following cases are worth notice as presenting unusual points of variation. According to Godron (3/101. 'De l'Espece' tome 1 page 406. Mr. Clark also refers to differences in the shape of the mammae. Godron states that in the Nubian race the scrotum is divided into two lobes; and Mr. Clark gives a ludicrous proof of this fact, for he saw in the Mauritius a male goat of the Muscat breed purchased at a high price for a female in full milk. These differences in the scrotum are probably not due to descent from distinct species: for Mr. Clark states that this part varies much in form.), the mammae differ greatly in shape in different breeds, being elongated in the common goat, hemispherical in the Angora race, and bilobed and divergent in the goats of Syria and Nubia. According to this same author, the males of certain breeds have lost their usual offensive odour. In one of the Indian breeds the males and females have horns of widely-different shapes (3/102. Mr. Clark 'Annals and Mag. of Nat. Hist.' 2nd series volume 2 1848 page 361.); and in some breeds the females are destitute of horns. (3/103. Desmarest 'Encyclop. Method. Mammalogie' page 480.) M. Ramu of Nancy informs me that many of the goats there bear on the upper part of the throat a pair of hairy appendages, 70 mm. in length and about 10 mm. in diameter, which in external appearance resemble those above described on the jaws of pigs. The presence of inter-digital pits or glands on all four feet has been thought to characterise the genus Ovis, and their absence to be characteristic of the genus Capra; but Mr. Hodgson has found that they exist in the front feet of the majority of Himalayan goats. (3/104. 'Journal of Asiatic Soc. of Bengal' volume 16 1847 pages 1020, 1025.) Mr. Hodgson measured the intestines in two goats of the Dugu race, and he found that the proportional length of the great and small intestines differed considerably. In one of these goats the caecum was thirteen inches, and in the other no less than thirty-six inches in length!

CHAPTER 1.IV

DOMESTIC RABBITS.

DOMESTIC RABBITS DESCENDED FROM THE COMMON WILD RABBIT. ANCIENT DOMESTICATION. ANCIENT SELECTION. LARGE LOP-EARED RABBITS. VARIOUS BREEDS. FLUCTUATING CHARACTERS. ORIGIN OF THE HIMALAYAN BREED. CURIOUS CASE OF INHERITANCE. FERAL RABBITS IN JAMAICA AND THE FALKLAND ISLANDS. PORTO SANTO FERAL RABBITS. OSTEOLOGICAL CHARACTERS. SKULL. SKULL OF HALF-LOP RABBITS. VARIATIONS IN THE SKULL ANALOGOUS TO DIFFERENCES IN DIFFERENT SPECIES OF HARES. VERTEBRAE. STERNUM. SCAPULA. EFFECTS OF USE AND DISUSE ON THE PROPORTIONS OF THE LIMBS AND BODY. CAPACITY OF THE SKULL AND REDUCED SIZE OF THE BRAIN. SUMMARY ON THE MODIFICATIONS OF DOMESTICATED RABBITS.

All naturalists, with, as far as I know, a single exception, believe that the several domestic breeds of the rabbit are descended from the common wild species; I shall therefore describe them more carefully than in the previous cases. Professor Gervais (4/1. M.P. Gervais 'Hist. Nat. des Mammiferes' 1854. tome 1 page 288.) states "that the true wild rabbit is smaller than the domestic; its proportions are not absolutely the same; its tail is smaller; its ears are shorter and more thickly clothed with hair; and these characters, without speaking of colour, are so many indications opposed to the opinion which unites these animals under the same specific denomination." Few naturalists will agree with this author that such slight differences are sufficient to separate as distinct species the wild and domestic rabbit. How extraordinary it would be, if close confinement, perfect tameness, unnatural food, and careful breeding, all prolonged during many generations, had not produced at least some effect! The tame rabbit has been domesticated from an ancient period. Confucius ranges rabbits among animals worthy to be sacrificed to the gods, and, as he prescribes their multiplication, they were probably at this early period domesticated in China. They are mentioned by several of the classical writers. In 1631 Gervaise Markham writes, "You shall not, as in other cattell, looke to their shape, but to their richnesse, onely elect your buckes, the largest and goodliest conies you can get; and for the richnesse of the skin, that is accounted the richest which hath the equallest mixture of blacke and white haire together, yet the blacke rather shadowing the white; the furre should be thicke, deepe, smooth, and shining;...they are of body much fatter and larger, and, when another skin is worth two or three pence, they are worth two shillings." From this full description we see that silver-grey rabbits existed in England at this period; and what is far more important, we see that the breeding or selection of rabbits was then carefully attended to. Aldrovandi, in 1637, describes, on the authority of several old writers (as Scaliger, in 1557), rabbits of various colours, some "like a hare," and he adds that P. Valerianus (who died a very old man in 1558) saw at Verona rabbits four times bigger than ours. (4/2. U. Aldrovandi 'De Quadrupedibus digitatis' 1637 page 383. For Confucius and G. Markham see a writer who has studied the subject in 'Cottage Gardener' January 22, 1861 page 250.)

From the fact of the rabbit having been domesticated at an ancient period, we must look to the northern hemisphere of the Old World, and to the warmer temperate regions alone, for the aboriginal parent-form; for the rabbit cannot live without protection in countries as cold as Sweden, and, though it has run wild in the tropical island of Jamaica, it has never greatly multiplied there. It now exists, and has long existed, in the warmer temperate parts of Europe, for fossil remains have been found in several countries. (4/3. Owen 'British Fossil Mammals' page 212.) The domestic rabbit readily becomes feral in these same countries, and when variously coloured kinds are turned out they generally revert to the ordinary grey colour. (4/4. Bechstein 'Naturgesch. Deutschlands' 1801 b. 1 page 1133. I have received similar accounts with respect to England and Scotland.) Wild rabbits, if taken young, can be domesticated, though the process is generally very troublesome. (4/5. 'Pigeons and Rabbits' by E.S. Delamer 1854 page 133. Sir J. Sebright 'Observations on Instinct' 1836 page 10) speaks most strongly on the difficulty. But this difficulty is not invariable, as I have received two accounts of perfect success in taming and breeding from the wild rabbit. See also Dr. P. Broca in 'Journal de la Physiologie' tome 2 page 368.) The various domestic races are often crossed, and are believed to be quite fertile together, and a perfect gradation can be shown to exist from the largest domestic kinds, having enormously developed ears, to the common wild kind. The parent-form must have been a burrowing animal, a habit not common, as far as I can discover, to any other species in the large genus Lepus. Only one wild species is known with certainty to exist in Europe; but the rabbit (if it be a true rabbit) from Mount Sinai, and likewise that from Algeria, present slight differences; and these forms have been considered by some authors as specifically distinct. (4/6. Gervais 'Hist. Nat. des Mammiferes' tome 1 page 292.) But such slight differences would aid us little in explaining the more considerable differences characteristic of the several domestic races. If the latter are the descendants of two or more closely allied species, these, with the exception of the common rabbit, have been exterminated in a wild state; and this is very improbable, seeing with what pertinacity this animal holds its ground. From these several reasons we may infer with safety that all the domestic breeds are the descendants of the common wild species. But from what we hear of the marvellous success in France in rearing hybrids between the hare and rabbit (4/7. See Dr. P. Broca's interesting memoir on this subject in Brown-Sequard 'Journ. de. Phys.' volume 2 page 367.), it is possible, though not probable, from the great difficulty in making the first cross, that some of the larger races, which are coloured like the hare, may have been modified by crosses with this animal. Nevertheless, the chief differences in the skeletons of the several domestic breeds cannot, as we shall presently see, have been derived from a cross with the hare.

There are many breeds which transmit their characters more or less truly. Every one has seen the enormous lop-eared rabbits exhibited at our shows; various allied sub-breeds are reared on the Continent, such as the so- called Andalusian, which is said to have a large head with a round forehead, and to attain a greater size than any other kind; another large Paris breed is named the Rouennais, and has a square head; the so-called Patagonian rabbit has remarkably short ears and a large round head. Although I have not seen all these breeds, I feel some doubt about there being any marked difference in the shape of their skulls. (4/8. The skulls of these breeds are briefly described in the 'Journal of Horticulture' May 7, 1861 page 108.) English lop-eared rabbits often weigh 8 pounds or 10 pounds, and one has been exhibited weighing 18 pounds; whereas a full-sized wild rabbit weighs only about 3 1/4 pounds. The head or skull in all the large lop-eared rabbits examined by me is much longer relatively to its breadth than in the wild rabbit. Many of them have loose transverse folds of skin or dewlaps beneath the throat, which can be pulled out so as to reach nearly to the ends of the jaws. Their ears are prodigiously developed, and hang down on each side of their faces. A rabbit was exhibited in 1867 with its two ears, measured from the tip of one to the tip of the other, 22 inches in length, and each ear 5 3/8 inches in breadth. In 1869 one was exhibited with ears, measured in the same manner, 23 1/8 in length and 5 1/2 in breadth; "thus exceeding any rabbit ever exhibited at a prize show." In a common wild rabbit I found that the length of two ears, from tip to tip, was 7 5/8 inches, and the breadth only 1 7/8 inch. The weight of body in the larger rabbits, and the development of their ears, are the qualities which win prizes, and have been carefully selected.

The hare-coloured, or, as it is sometimes called, the Belgian rabbit, differs in nothing except colour from the other large breeds; but Mr. J. Young, of Southampton, a great breeder of this kind, informs me that the females, in all the specimens examined by him, had only six mammae; and this certainly was the case with two females which came into my possession. Mr. B.P. Brent, however, assures me that the number is variable with other domestic rabbits. The common wild rabbit always has ten mammae. The Angora rabbit is remarkable from the length and fineness of its fur, which even on the soles of the feet is of considerable length. This breed is the only one which differs in its mental qualities, for it is said to be much more sociable than other rabbits, and the male shows no wish to destroy its young. (4/9. 'Journal of Horticulture' 1861 page 380.) Two live rabbits were brought to me from Moscow, of about the size of the wild species, but with long soft fur, different from that of the Angora. These Moscow rabbits had pink eyes and were snow-white, excepting the ears, two spots near the nose, the upper and under surface of the tail, and the hinder tarsi, which were blackish-brown. In short, they were coloured nearly like the so-called Himalayan rabbits, presently to be described, and differed from them only in the character of their fur. There are two other breeds which come true to colour, but differ in no other respect, namely silver-greys and chinchillas. Lastly, the Nicard or Dutch rabbit may be mentioned, which varies in colour, and is remarkable from its small size, some specimens weighing only 1 1/4 pounds; rabbits of this breed make excellent nurses for other and more delicate kinds. (4/10. 'Journal of Horticulture' May 28, 1861 page 169.)

(FIGURE 5. HALF-LOP RABBIT. (Copied from E.S. Delamer's work.)

Certain characters are remarkably fluctuating, or are very feebly transmitted by domestic rabbits: thus, one breeder tells me that with the smaller kinds he has hardly ever raised a whole litter of the same colour: with the large lop-eared breeds "it is impossible," says a great judge (4/11. 'Journal of Horticulture' 1861 page 327. With respect to the ears see Delamer on 'Pigeons and Rabbits' 1854 page 141; also 'Poultry Chronicle' volume 2 page 499 and ditto for 1854 page 586.), "to breed true to colour, but by judicious crossing a great deal may be done towards it. The fancier should know how his does are bred, that is, the colour of their parents." Nevertheless, certain colours, as we shall presently see, are transmitted truly. The dewlap is not strictly inherited. Lop-eared rabbits, with their ears hanging down flat on each side of the face, do not transmit this character at all truly. Mr. Delamer remarks that, "with fancy rabbits, when both the parents are perfectly formed, have model ears, and are handsomely marked, their progeny do not invariably turn out the same." When one parent, or even both, are oar-laps, that is, have their ears sticking out at right angles, or when one parent or both are half-lops, that is, have only one ear dependent, there is nearly as good a chance of the progeny having both ears full-lop, as if both parents had been thus characterised. But I am informed, if both parents have upright ears, there is hardly a chance of a full-lop. In some half-lops the ear that hangs down is broader and longer than the upright ear (4/12. Delamer 'Pigeons and Rabbits' page 136. See also 'Journal of Horticulture' 1861 page 375.); so that we have the unusual case of a want of symmetry on the two sides. This difference in the position and size of the two ears probably indicates that the lopping results from the great length and weight of the ear, favoured no doubt by the weakness of the muscles consequent on disuse. Anderson (4/13. 'An Account of the different Kinds of Sheep in the Russian Dominions' 1794 page 39.) mentions a breed having only a single ear; and Professor Gervais another breed destitute of ears.

We come now to the Himalayan breed, which is sometimes called Chinese, Polish, or Russian. These pretty rabbits are white, or occasionally yellow, excepting their ears, nose, feet, and the upper side of the tail, which are all brownish-black; but as they have red eyes, they may be considered as albinoes. I have received several accounts of their breeding perfectly true. From their symmetrical marks, they were at first ranked as specifically distinct, and were provisionally named L. nigripes. (4/14. 'Proc. Zoolog. Soc.' June 23, 1857 page 159.) Some good observers thought that they could detect a difference in their habits, and stoutly maintained that they formed a new species. The origin of this breed is so curious, both in itself and as throwing some light on the complex laws of inheritance that it is worth giving in detail. But it is first necessary briefly to describe two other breeds: silver-greys or silver-sprigs generally have black heads and legs, and their fine grey fur is interspersed with numerous black and white long hairs. They breed perfectly true, and have long been kept in warrens. When they escape and cross with common rabbits, the product, as I hear from Mr. Wyrley Birch, of Wretham Hall, is not a mixture of the two colours, but about half take after the one parent, and the other half after the other parent. Secondly, chinchillas or tame silver-greys (I will use the former name) have short, paler, mouse or slate-coloured fur, interspersed with long, blackish, slate-coloured, and white hairs. (4/15. 'Journal of Horticulture' April 9, 1861 page 35.) These rabbits breed perfectly true. A writer stated in 1857 (4/16. 'Cottage Gardener' 1857 page 141.) that he had produced Himalayan rabbits in the following manner. He had a breed of chinchillas which had been crossed with the common black rabbit, and their offspring were either blacks or chinchillas. These latter were again crossed with other chinchillas (which had also been crossed with silver-greys), and from this complicated cross Himalayan rabbits were raised. From these and other similar statements, Mr. Bartlett (4/17. Mr. Bartlett in 'Proc. Zoolog Soc.' 1861 page 40.) was led to make a careful trial in the Zoological Gardens, and he found that by simply crossing silver-greys with chinchillas he could always produce some few Himalayans; and the latter, notwithstanding their sudden origin, if kept separate, bred perfectly true. But I have recently been assured the pure silver-greys of any sub-breed occasionally produce Himalayans.

The Himalayans, when first born, are quite white, and are then true albinoes; but in the course of a few months they gradually assume their dark ears, nose, feet, and tail. Occasionally, however, as I am informed by Mr. W.A. Wooler and the Rev. W.D. Fox, the young are born of a very pale grey colour, and specimens of such fur were sent me by the former gentleman. The grey tint, however, disappears as the animal comes to maturity. So that with these Himalayans there is a tendency, strictly confined to early youth, to revert to the colour of the adult silver-grey parent-stock. Silver-greys and chinchillas, on the other hand, present a remarkable contrast with the Himalayans in their colour whilst quite young, for they are born perfectly black, but soon assume their characteristic grey or silver tints. The same thing occurs with grey horses, which, as long as they are foals, are generally of a nearly black colour, but soon become grey, and get whiter and whiter as they grow older. Hence the usual rule is that Himalayans are born white and afterwards become in certain parts of their bodies dark-coloured; whilst silver-greys are born black and afterwards become sprinkled with white. Exceptions, however, and of a directly opposite nature, occasionally occur in both cases. For young silver-greys are sometimes born in warrens, as I hear from Mr. W. Birch, of a cream-colour, but these young animals ultimately become black. The Himalayans, on the other hand, sometimes produce, as is stated by an experienced amateur (4/18. 'Phenomenon in Himalayan Rabbits' in 'Journal of Horticulture' January 27, 1865 page 102.), a single black young one in a litter; and this, before two months elapse, becomes perfectly white.

To sum up the whole curious case: wild silver-greys may be considered as black rabbits which become grey at an early period of life. When they are crossed with common rabbits, the offspring are said not to have blended colours, but to take after either parent; and in this respect they resemble black and albino varieties of most quadrupeds, which often transmit their colours in this same manner. When they are crossed with chinchillas, that is, with a paler sub-variety, the young are at first pure albinoes, but soon become dark-coloured in certain parts of their bodies, and are then called Himalayans. The young Himalayans, however, are sometimes at first either pale grey or completely black, in either case changing after a time to white. In a future chapter I shall advance a large body of facts showing that, when two varieties are crossed both of which differ in colour from their parent-stock, there is a strong tendency in the young to revert to the aboriginal colour; and what is very remarkable, this reversion occasionally supervenes, not before birth, but during the growth of the animal. Hence, if it could be shown that silver-greys and chinchillas were the offspring of a cross between a black and albino variety with the colours intimately blended — a supposition in itself not improbable, and supported by the circumstance of silver-greys in warrens sometimes producing creamy-white young, which ultimately become black — then all the above given paradoxical facts on the changes of colour in silver-greys and in their descendants the Himalayans would come under the law of reversion, supervening at different periods of growth and in different degrees, either to the original black or to the original albino parent-variety.

It is, also, remarkable that Himalayans, though produced so suddenly; breed true. But as, whilst young, they are albinoes, the case falls under a very general rule; albinism being well known to be strongly inherited, for instance with white mice and many other quadrupeds, and even white flowers. But why, it may be asked, do the ears, tail, nose, and feet, and no other part of the body, revert to a black colour? This apparently depends on a law, which generally holds good, namely, that characters common to many species of a genus — and this, in fact, implies long inheritance from the ancient progenitor of the genus — are found to resist variation, or to reappear if lost, more persistently than the characters which are confined to the separate species. Now, in the genus Lepus, a large majority of the species have their ears and the upper surface of the tail tinted black; but the persistence of these marks is best seen in those species which in winter become white: thus, in Scotland the L. variabilis (4/19. G.R. Waterhouse 'Natural History of Mammalia: Rodents' 1846 pages 52, 60, 105.) in its winter dress has a shade of colour on its nose, and the tips of its ears are black: in the L. tibetanus the ears are black, the upper surface of the tail greyish-black, and the soles of the feet brown: in L. glacialis the winter fur is pure white, except the soles of the feet and the points of the ears. Even in the variously-coloured fancy rabbits we may often observe a tendency in these same parts to be more darkly tinted than the rest of the body. Thus the several coloured marks on the Himalayan rabbits, as they grow old, are rendered intelligible. I may add a nearly analogous case: fancy rabbits very often have a white star on their foreheads; and the common English hare, whilst young, generally has, as I have myself observed, a similar white star on its forehead.

When variously coloured rabbits are set free in Europe, and are thus placed under their natural conditions, they generally revert to the aboriginal grey colour; this may be in part due to the tendency in all crossed animals, as lately observed, to revert to their primordial state. But this tendency does not always prevail; thus silver-grey rabbits are kept in warrens, and remain true though living almost in a state of nature; but a warren must not be stocked with both silver-greys and common rabbits; otherwise "in a few years there will be none but common greys surviving." (4/20. Delamer on 'Pigeons and Rabbits' page 114.) When rabbits run wild in foreign countries under new conditions of life, they by no means always revert to their aboriginal colour. In Jamaica the feral rabbits are described as having been "slate-coloured, deeply tinted with sprinklings of white on the neck, on the shoulders, and on the back; softening off to blue-white under the breast and belly." (4/21. Gosse 'Sojourn in Jamaica' 1851 page 441 as described by an excellent observer, Mr. R. Hill. This is the only known case in which rabbits have become feral in a hot country. They can be kept, however, at Loanda (see Livingstone 'Travels' page 407). In parts of India, as I am informed by Mr. Blyth, they breed well.) But in this tropical island the conditions were not favourable to their increase, and they never spread widely, and are now extinct, as I hear from Mr. R. Hill, owing to a great fire which occurred in the woods. Rabbits during many years have run wild in the Falkland Islands; they are abundant in certain parts, but do not spread extensively. Most of them are of the common grey colour; a few, as I am informed by Admiral Sulivan, are hare- coloured, and many are black, often with nearly symmetrical white marks on their faces. Hence, M. Lesson described the black variety as a distinct species, under the name of Lepus magellanicus, but this, as I have elsewhere shown, is an error. (4/22. Darwin 'Journal of Researches' page 193; and 'Zoology of the Voyage of the Beagle: Mammalia' page 92.) Within recent times the sealers have stocked some of the small outlying islets in the Falkland group with rabbits; and on Pebble Islet, as I hear from Admiral Sulivan, a large proportion are hare-coloured, whereas on Rabbit Islet a large proportion are of a bluish colour, which is not elsewhere seen. How the rabbits were coloured which were turned out of these islets is not known.

The rabbits which have become feral on the island of Porto Santo, near Madeira, deserve a fuller account. In 1418 or 1419, J. Gonzales Zarco (4/23. Kerr 'Collection of Voyages' volume 2 page 177: page 205 for Cada Mosto. According to a work published in Lisbon in 1717 entitled 'Historia Insulana' written by a Jesuit, the rabbits were turned out in 1420. Some authors believe that the island was discovered in 1413.) happened to have a female rabbit on board which had produced young during the voyage, and he turned them all out on the island. These animals soon increased so rapidly, that they became a nuisance, and actually caused the abandonment of the settlement. Thirty-seven years subsequently, Cada Mosto describes them as innumerable; nor is this surprising, as the island was not inhabited by any beast of prey or by any terrestrial mammal. We do not know the character of the mother-rabbit; but it was probably the common domesticated kind. The Spanish peninsula, whence Zarco sailed, is known to have abounded with the common wild species at the most remote historical period; and as these rabbits were taken on board for food, it is improbable that they should have been of any peculiar breed. That the breed was well domesticated is shown by the doe having littered during the voyage. Mr. Wollaston, at my request, brought home two of these feral rabbits in spirits of wine; and, subsequently, Mr. W. Haywood sent to me three more specimens in brine, and two alive. These seven specimens, though caught at different periods, closely resembled each other. They were full grown, as shown by the state of their bones. Although the conditions of life in Porto Santo are evidently highly favourable to rabbits, as proved by their extraordinarily rapid increase, yet they differ conspicuously in their small size from the wild English rabbit. Four English rabbits, measured from the incisors to the anus, varied between 17 and 17 3/4 inches in length; whilst two of the Porto Santo rabbits were only 14 1/2 and 15 inches in length. But the decrease in size is best shown by weight; four wild English rabbits averaged 3 pounds 5 ounces, whilst one of the Porto Santo rabbits, which had lived for four years in the Zoological Gardens, but had become thin, weighed only 1 pound 9 ounces. A fairer test is afforded by the comparison of the well-cleaned limb-bones of a Porto Santo rabbit killed on the island with the same bones of a wild English rabbit of average size, and they differed in the proportion of rather less than five to nine. So that the Porto Santo rabbits have decreased nearly three inches in length, and almost half in weight of body. (4/24. Something of the same kind has occurred on the island of Lipari, where, according to Spallanzani ('Voyage dans les deux Siciles' quoted by Godron 'De l'Espece' page 364), a countryman turned out some rabbits which multiplied prodigiously, but, says Spallanzani, "les lapins de l'ile de Lipari sont plus petits que ceux qu'on eleve en domesticite.") The head has not decreased in length proportionally with the body; and the capacity of the brain case is, as we shall hereafter see, singularly variable. I prepared four skulls, and these resembled each other more closely than do generally the skulls of wild English rabbits; but the only difference in structure which they presented was that the supra-orbital processes of the frontal bones were narrower.

In colour the Porto Santo rabbit differs considerably from the common rabbit; the upper surface is redder, and is rarely interspersed with any black or black-tipped hairs. The throat and certain parts of the under surface, instead of being pure white, are generally pale grey or leaden colour. But the most remarkable difference is in the ears and tail; I have examined many fresh English rabbits, and the large collection of skins in the British Museum from various countries, and all have the upper surface of the tail and the tips of the ears clothed with blackish-grey fur; and this is given in most works as one of the specific characters of the rabbit. Now in the seven Porto Santo rabbits the upper surface of the tail was reddish-brown, and the tips of the ears had no trace of the black edging. But here we meet with a singular circumstance: in June, 1861 I examined two of these rabbits recently sent to the Zoological Gardens, and their tails and ears were coloured as just described; but when one of their dead bodies was sent to me in February, 1865, the ears were plainly edged, and the upper surface of the tail was covered with blackish-grey fur, and the whole body was much less red; so that under the English climate this individual rabbit had recovered the proper colour of its fur in rather less than four years!

The two little Porto Santo rabbits, whilst alive in the Zoological Gardens, had a remarkably different appearance from the common kind. They were extraordinarily wild and active, so that many persons exclaimed on seeing them that they were more like large rats than rabbits. They were nocturnal to an unusual degree in their habits, and their wildness was never in the least subdued; so that the superintendent, Mr. Bartlett, assured me that he had never had a wilder animal under his charge. This is a singular fact, considering that they are descended from a domesticated breed. I was so much surprised at it, that I requested Mr. Haywood to make inquiries on the spot, whether they were much hunted by the inhabitants, or persecuted by hawks, or cats, or other animals; but this is not the case, and no cause can be assigned for their wildness. They live both on the central, higher rocky land and near the sea-cliffs, and, from being exceedingly shy and timid, seldom appear in the lower and cultivated districts. They are said to produce from four to six young at a birth, and their breeding season is in July and August. Lastly, and this is a highly remarkable fact, Mr. Bartlett could never succeed in getting these two rabbits, which were both males, to associate or breed with the females of several breeds which were repeatedly placed with them.

If the history of these Porto Santo rabbits had not been known, most naturalists, on observing their much reduced size, their colour, reddish above and grey beneath, their tails and ears not tipped with black, would have ranked them as a distinct species. They would have been strongly confirmed in this view by seeing them alive in the Zoological Gardens, and hearing that they refused to couple with other rabbits. Yet this rabbit, which there can be little doubt would thus have been ranked as a distinct species, as certainly originated since the year 1420. Finally, from the three cases of the rabbits which have run wild in Porto Santo, Jamaica, and the Falkland Islands, we see that these animals do not, under new conditions of life, revert to or retain their aboriginal character, as is so generally asserted to be the case by most authors.

OSTEOLOGICAL CHARACTERS.

When we remember, on the one hand, how frequently it is stated that important parts of the structure never vary; and, on the other hand, on what small differences in the skeleton fossil species have often been founded, the variability of the skull and of some other bones in the domesticated rabbit well deserves attention. It must not be supposed that the more important differences immediately to be described strictly characterise any one breed; all that can be said is, that they are generally present in certain breeds. We should bear in mind that selection has not been applied to fix any character in the skeleton, and that the animals have not had to support themselves under uniform habits of life. We cannot account for most of the differences in the skeleton; but we shall see that the increased size of the body, due to careful nurture and continued selection, has affected the head in a particular manner. Even the elongation and lopping of the ears have influenced in a small degree the form of the whole skull. The want of exercise has apparently modified the proportional length of the limbs in comparison with that of the body.