The Variation of Animals and Plants under Domestication — Volume 1

There is another and closely allied difficulty consequent on the doctrine of the descent of our domestic dogs from several wild species, namely, that they do not seem to be perfectly fertile with their supposed parents. But the experiment has not been quite fairly tried; the Hungarian dog, for instance, which in external appearance so closely resembles the European wolf, ought to be crossed with this wolf: and the pariah dogs of India with Indian wolves and jackals; and so in other cases. That the sterility is very slight between certain dogs and wolves and other Canidae is shown by savages taking the trouble to cross them. Buffon got four successive generations from the wolf and dog, and the mongrels were perfectly fertile together. (1/48. M. Broca has shown ('Journal de Physiologie' tome 2 page 353) that Buffon's experiments have been often misrepresented. Broca has collected (pages 390-395) many facts on the fertility of crossed dogs, wolves, and jackals.) But more lately M. Flourens states positively as the result of his numerous experiments that hybrids from the wolf and dog, crossed inter se, become sterile at the third generation, and those from the jackal and dog at the fourth generation. (1/49. 'De la Longevite Humaine' par M. Flourens 1855 page 143. Mr. Blyth says ('Indian Sporting Review' volume 2 page 137) that he has seen in India several hybrids from the pariah-dog and jackal; and between one of these hybrids and a terrier. The experiments of Hunter on the jackal are well-known. See also Isid. Geoffroy St. — Hilaire, 'Hist. Nat. Gen.' tome 3 page 217, who speaks of the hybrid offspring of the jackal as perfectly fertile for three generations.) But these animals were closely confined; and many wild animals, as we shall see in a future chapter, are rendered by confinement in some degree or even utterly sterile. The Dingo, which breeds freely in Australia with our imported dogs, would not breed though repeatedly crossed in the Jardin des Plantes. (1/50. On authority of F. Cuvier quoted in Bronn's 'Geschichte der Natur' b. 2 s. 164.) Some hounds from Central Africa, brought home by Major Denham, never bred in the Town of London (1/51. W.C.L. Martin 'History of the Dog' 1845 page 203. Mr. Philip P. King, after ample opportunities of observation, informs me that the Dingo and European dogs often cross in Australia.); and a similar tendency to sterility might be transmitted to the hybrid offspring of a wild animal. Moreover, it appears that in M. Flourens' experiments the hybrids were closely bred in and in for three or four generations; and this circumstance would most certainly increase the tendency to sterility. Several years ago I saw confined in the Zoological Gardens of London a female hybrid from an English dog and jackal, which even in this the first generation was so sterile that, as I was assured by her keeper, she did not fully exhibit her proper periods; but this case was certainly exceptional, as numerous instances have occurred of fertile hybrids from these two animals. In almost all experiments on the crossing of animals there are so many causes of doubt, that it is extremely difficult to come to any positive conclusion. It would, however, appear, that those who believe that our dogs are descended from several species will have not only to admit that their offspring after a long course of domestication generally lose all tendency to sterility when crossed together; but that between certain breeds of dogs and some of their supposed aboriginal parents a certain degree of sterility has been retained or possibly even acquired.

Notwithstanding the difficulties in regard to fertility given in the last two paragraphs, when we reflect on the inherent improbability of man having domesticated throughout the world one single species alone of so widely distributed, so easily tamed, and so useful a group as the Canidae; when we reflect on the extreme antiquity of the different breeds; and especially when we reflect on the close similarity, both in external structure and habits, between the domestic dogs of various countries and the wild species still inhabiting these same countries, the balance of evidence is strongly in favour of the multiple origin of our dogs.

DIFFERENCES BETWEEN THE SEVERAL BREEDS OF THE DOG.

If the several breeds have descended from several wild stocks, their difference can obviously in part be explained by that of their parent species. For instance, the form of the greyhound may be partly accounted for by descent from some such animal as the slim Abyssinian Canis simensis (1/52. Ruppel 'Neue Wirbelthiere von Abyssinien' 1835-40 'Mammif.' s. 39 pl. 14. There is a specimen of this fine animal in the British Museum.), with its elongated muzzle; that of the larger dogs from the larger wolves, and the smaller and slighter dogs from the jackals: and thus perhaps we may account for certain constitutional and climatal differences. But it would be a great error to suppose that there has not been in addition (1/53. Even Pallas admits this; see 'Act. Acad. St. Petersburgh' 1780 page 93.) a large amount of variation. The intercrossing of the several aboriginal wild stocks, and of the subsequently formed races, has probably increased the total number of breeds, and, as we shall presently see, has greatly modified some of them. But we cannot explain by crossing the origin of such extreme forms as thoroughbred greyhounds, bloodhounds, bulldogs, Blenheim spaniels, terriers, pugs, etc., unless we believe that forms equally or more strongly characterised in these different respects once existed in nature. But hardly any one has been bold enough to suppose that such unnatural forms ever did or could exist in a wild state. When compared with all known members of the family of Canidae they betray a distinct and abnormal origin. No instance is on record of such dogs as bloodhounds, spaniels, true greyhounds having been kept by savages: they are the product of long-continued civilisation.

[The number of breeds and sub-breeds of the dog is great; Youatt for instance, describes twelve kinds of greyhounds. I will not attempt to enumerate or describe the varieties, for we cannot discriminate how much of their difference is due to variation, and how much to descent from different aboriginal stocks. But it may be worth while briefly to mention some points. Commencing with the skull, Cuvier has admitted (1/54. Quoted by I. Geoffroy 'Hist. Nat. Gen.' tome 3 page 453.) that in form the differences are "plus fortes que celles d'aucunes especes sauvages d'un meme genre naturel." The proportions of the different bones; the curvature of the lower jaw, the position of the condyles with respect to the plane of the teeth (on which F. Cuvier founded his classification), and in mastiffs the shape of its posterior branch; the shape of the zygomatic arch, and of the temporal fossae; the position of the occiput — all vary considerably. (1/55. F. Cuvier in 'Annales du Museum' tome 18 page 337; Godron 'De l'Espece' tome 1 page 342; and Col. H. Smith in 'Nat. Library' volume 9 page 101. See also some observations on the degeneracy of the skull in certain breeds, by Prof. Bianconi 'La Theorie Darwinienne' 1874 page 279.) The difference in size between the brains of dogs belonging to large and small breeds "is something prodigious." "Some dogs' brains are high and rounded, while others are low, long, and narrow in front." In the latter, "the olfactory lobes are visible for about half their extent, when the brain is seen from above, but they are wholly concealed by the hemispheres in other breeds." (1/56. Dr. Burt Wilder 'American Assoc. Advancement of Science' 1873 pages 236, 239.) The dog has properly six pairs of molar teeth in the upper jaw, and seven in the lower; but several naturalists have seen not rarely an additional pair in the upper jaw (1/57. Isid. Geoffroy Saint-Hilaire 'Hist. des Anomalies' 1832 tome 1 page 660, Gervais 'Hist. Nat. des Mammiferes' tome 2 1855 page 66. De Blainville ('Osteographie, Canidae' page 137) has also seen an extra molar on both sides.); and Professor Gervais says that there are dogs "qui ont sept paires de dents superieures et huit inferieures." De Blainville (1/58. 'Osteographie, Canidae' page 137.) has given full particulars on the frequency of these deviations in the number of the teeth, and has shown that it is not always the same tooth which is supernumerary. In short- muzzled races, according to H. Muller (1/59. Wurzburger 'Medecin. Zeitschrift' 1860 b. 1 s. 265.), the molar teeth stand obliquely, whilst in long-muzzled races they are placed longitudinally, with open spaces between them. The naked, so-called Egyptian or Turkish dog is extremely deficient in its teeth (1/60. Mr. Yarrell in 'Proc. Zoological Soc.' October 8, 1833. Mr. Waterhouse showed me a skull of one of these dogs, which had only a single molar on each side and some imperfect incisors.), — sometimes having none except one molar on each side; but this, though characteristic of the breed, must be considered as a monstrosity. M. Girard (1/61. Quoted in 'The Veterinary' London volume 8 page 415.), who seems to have attended closely to the subject, says that the period of the appearance of the permanent teeth differs in different dogs, being earlier in large dogs; thus the mastiff assumes its adult teeth in four or five months, whilst in the spaniel the period is sometimes more than seven or eight months. On the other hand small dogs are mature, and the females have arrived at the best age for breeding, when one year old, whereas large dogs "are still in their puppyhood at this time, and take fully twice as long to develop their proportions." (1/62. This is quoted from Stonehenge, a great authority, 'The Dog' 1867 page 187.)

With respect to minor differences little need be said. Isidore Geoffroy has shown (1/63. 'Hist. Nat. General' tome 3 page 448.) that in size some dogs are six times as long (the tail being excluded) as others; and that the height relatively to the length of the body varies from between one to two, and one to nearly four. In the Scotch deer-hound there is a striking and remarkable difference in the size of the male and female. (1/64. W. Scrope 'Art of Deer-Stalking' page 354.) Every one knows how the ears vary in size in different breeds, and with their great development their muscles become atrophied. Certain breeds of dogs are described as having a deep furrow between the nostrils and lips. The caudal vertebrae, according to F. Cuvier, on whose authority the two last statements rest, vary in number; and the tail in English cattle and some shepherd dogs is almost absent. The mammae vary from seven to ten in number; Daubenton, having examined twenty- one dogs, found eight with five mammae on each side; eight with four on each side; and the others with an unequal number on the two sides. (1/65. Quoted by Col. Ham. Smith in 'Nat. Lib.' volume 10 page 79.) Dogs have properly five toes in front and four behind, but a fifth toe is often added; and F. Cuvier states that, when a fifth toe is present, a fourth cuneiform bone is developed; and, in this case, sometimes the great cuneiform bone is raised, and gives on its inner side a large articular surface to the astragalus; so that even the relative connection of the bones, the most constant of all characters, varies. These modifications, however, in the feet of dogs are not important, because they ought to be ranked, as De Blainville has shown (1/66. De Blainville 'Osteographie, Canidae' page 134. F. Cuvier 'Annales du Museum' tome 18 page 342. In regard to mastiffs, see Col. H. Smith 'Nat. Lib.' volume 10 page 218. For the Thibet mastiff, see Mr. Hodgson in 'Journal of As. Soc. of Bengal' volume 1 1832 page 342.) as monstrosities. Nevertheless they are interesting from being correlated with the size of the body, for they occur much more frequently with mastiffs and other large breeds than with small dogs. Closely allied varieties, however, sometimes differ in this respect; thus Mr. Hodgson states that the black-and-tan Lassa variety of the Thibet mastiff has the fifth digit, whilst the Mustang sub-variety is not thus characterised. The extent to which the skin is developed between the toes varies much; but we shall return to this point. The degree to which the various breeds differ in the perfection of their senses, dispositions, and inherited habits is notorious to every one. The breeds present some constitutional differences: the pulse, says Youatt (1/67. 'The Dog' 1845 page 186. With respect to diseases Youatt asserts (page 167) that the Italian greyhound is "strongly subject" to polypi in the matrix or vagina. The spaniel and pug (page 182) are most liable to bronchocele. The liability to distemper (page 232) is extremely different in different breeds. On the distemper, see also Col. Hutchinson on 'Dog Breaking' 1850 page 279.) "varies materially according to the breed, as well as to the size of the animal." Different breeds of dogs are subject in different degrees to various diseases. They certainly become adapted to different climates under which they have long existed. It is notorious that most of our best European breeds deteriorate in India. (1/68. See 'Youatt on the Dog' page 15; 'The Veterinary' London volume 11 page 235.) The Rev R. Everest (1/69. 'Journal of As. Soc. of Bengal' volume 3 page 19.) believes that no one has succeeded in keeping the Newfoundland dog long alive in India; so it is, according to Lichtenstein (1/70. 'Travels' volume 2 page 15.), even at the Cape of Good Hope. The Thibet mastiff degenerates on the plains of India, and can live only on the mountains. (1/71. Hodgson in 'Journal of As. Soc. of Bengal' volume 1 page 342.) Lloyd (1/72. 'Field Sports of the North of Europe' volume 2 page 165.) asserts that our bloodhounds and bulldogs have been tried, and cannot withstand the cold of the northern European forests.]

Seeing in how many characters the races of the dog differ from each other, and remembering Cuvier's admission that their skulls differ more than do those of the species of any natural genus, and bearing in mind how closely the bones of wolves, jackals, foxes, and other Canidae agree, it is remarkable that we meet with the statement, repeated over and over again, that the races of the dog differ in no important characters. A highly competent judge, Prof. Gervais (1/73. 'Hist. Nat. des Mammif.' 1855 tome 2 pages 66, 67.), admits "si l'on prenait sans controle les alterations dont chacun de ces organes est susceptible, on pourrait croire qu'il y a entre les chiens domestiques des differences plus grandes que celles qui separent ailleurs les especes, quelquefois meme les genres." Some of the differences above enumerated are in one respect of comparatively little value, for they are not characteristic of distinct breeds: no one pretends that such is the case with the additional molar teeth or with the number of mammae; the additional digit is generally present with mastiffs, and some of the more important differences in the skull and lower jaw are more or less characteristic of various breeds. But we must not forget that the predominant power of selection has not been applied in any of these cases; we have variability in important parts, but the differences have not been fixed by selection. Man cares for the form and fleetness of his greyhounds, for the size of his mastiffs, and formerly for the strength of the jaw in his bulldogs, etc.; but he cares nothing about the number of their molar teeth or mammae or digits; nor do we know that differences in these organs are correlated with, or owe their development to, differences in other parts of the body about which man does care. Those who have attended to the subject of selection will admit that, nature having given variability, man, if he so chose, could fix five toes to the hinder feet of certain breeds of dogs, as certainly as to the feet of his Dorking fowls: he could probably fix, but with much more difficulty, an additional pair of molar teeth in either jaw, in the same way as he has given additional horns to certain breeds of sheep; if he wished to produce a toothless breed of dogs, having the so-called Turkish dog with its imperfect teeth to work on, he could probably do so, for he has succeeded in making hornless breeds of cattle and sheep.

With respect to the precise causes and steps by which the several races of dogs have come to differ so greatly from each other, we are, as in most other cases, profoundly ignorant. We may attribute part of the difference in external form and constitution to inheritance from distinct wild stocks, that is to changes effected under nature before domestication. We must attribute something to the crossing of the several domestic and natural races. I shall, however, soon recur to the crossing of races. We have already seen how often savages cross their dogs with wild native species; and Pennant gives a curious account (1/74. 'History of Quadrupeds' 1793 volume 1 page 238.) of the manner in which Fochabers, in Scotland, was stocked "with a multitude of curs of a most wolfish aspect" from a single hybrid-wolf brought into that district.

It would appear that climate to a certain extent directly modifies the forms of dogs. We have lately seen that several of our English breeds cannot live in India, and it is positively asserted that when bred there for a few generations they degenerate not only in their mental faculties, but in form. Captain Williamson (1/75. 'Oriental Field Sports' quoted by Youatt 'The Dog' page 15.), who carefully attended to this subject, states that "hounds are the most rapid in their decline;" "greyhounds and pointers, also, rapidly decline." But spaniels, after eight or nine generations, and without a cross from Europe, are as good as their ancestors. Dr. Falconer informs me that bulldogs, which have been known, when first brought into the country, to pin down even an elephant by its trunk, not only fall off after two or three generations in pluck and ferocity, but lose the under-hung character of their lower jaws; their muzzles become finer and their bodies lighter. English dogs imported into India are so valuable that probably due care has been taken to prevent their crossing with native dogs; so that the deterioration cannot be thus accounted for. The Rev. R. Everest informs me that he obtained a pair of setters, born in India, which perfectly resembled their Scotch parents: he raised several litters from them in Delhi, taking the most stringent precautions to prevent a cross, but he never succeeded, though this was only the second generation in India, in obtaining a single young dog like its parents in size or make; their nostrils were more contracted, their noses more pointed, their size inferior, and their limbs more slender. So again on the coast of Guinea, dogs, according to Bosman, "alter strangely; their ears grow long and stiff like those of foxes, to which colour they also incline, so that in three or four years, they degenerate into very ugly creatures; and in three or four broods their barking turns into a howl." (1/76. A. Murray gives this passage in his 'Geographical Distribution of Mammals' 4to 1866 page 8.) This remarkable tendency to rapid deterioration in European dogs subjected to the climate of India and Africa, may be largely accounted for by reversion to a primordial condition which many animals exhibit, as we shall hereafter see, when their constitutions are in any way disturbed.

Some of the peculiarities characteristic of the several breeds of the dog have probably arisen suddenly, and, though strictly inherited, may be called monstrosities; for instance, the shape of the legs and body in the turnspit of Europe and India; the shape of the head and the under-hanging jaw in the bull-and pug-dog, so alike in this one respect and so unlike in all others. A peculiarity suddenly arising, and therefore in one sense deserving to be called a monstrosity, may, however, be increased and fixed by man's selection. We can hardly doubt that long-continued training, as with the greyhound in coursing hares, as with water-dogs in swimming — and the want of exercise, in the case of lapdogs — must have produced some direct effect on their structure and instincts. But we shall immediately see that the most potent cause of change has probably been the selection, both methodical and unconscious, of slight individual differences, — the latter kind of selection resulting from the occasional preservation, during hundreds of generations, of those individual dogs which were the most useful to man for certain purposes and under certain conditions of life. In a future chapter on Selection I shall show that even barbarians attend closely to the qualities of their dogs. This unconscious selection by man would be aided by a kind of natural selection; for the dogs of savages have partly to gain their own subsistence: for instance, in Australia, as we hear from Mr. Nind (1/77. Quoted by Mr. Galton 'Domestication of Animals' page 13.), the dogs are sometimes compelled by want to leave their masters and provide for themselves; but in a few days they generally return. And we may infer that dogs of different shapes, sizes, and habits, would have the best chance of surviving under different circumstances, — on open sterile plains, where they have to run down their own prey, — on rocky coasts, where they have to feed on crabs and fish left in the tidal pools, as in the case of New Guinea and Tierra del Fuego. In this latter country, as I am informed by Mr. Bridges, the Catechist to the Mission, the dogs turn over the stones on the shore to catch the crustaceans which lie beneath, and they "are clever enough to knock off the shell-fish at a first blow;" for if this be not done, shell-fish are well-known to have an almost invincible power of adhesion.

It has already been remarked that dogs differ in the degree to which their feet are webbed. In dogs of the Newfoundland breed, which are eminently aquatic in their habits, the skin, according to Isidore Geoffroy (1/78. 'Hist. Nat. Gen.' tome 3 page 450.), extends to the third phalanges whilst in ordinary dogs it extends only to the second. In two Newfoundland dogs which I examined, when the toes were stretched apart and viewed on the under side, the skin extended in a nearly straight line between the outer margins of the balls of the toes; whereas, in two terriers of distinct sub- breeds, the skin viewed in the same manner was deeply scooped out. In Canada there is a dog which is peculiar to the country and common there, and this has "half-webbed feet and is fond of the water." (1/79. Mr. Greenhow on the Canadian Dog in Loudon's 'Mag. of Nat. Hist.' volume 6 1833 page 511.) English otter-hounds are said to have webbed feet: a friend examined for me the feet of two, in comparison with the feet of some harriers and bloodhounds; he found the skin variable in extent in all, but more developed in the otter-hounds than in the others. (1/80. See Mr. C.O. Groom-Napier on the webbing of the hind feet of Otterhounds in 'Land and Water' October 13, 1866 page 270.) As aquatic animals which belong to quite different orders have webbed feet, there can be no doubt that this structure would be serviceable to dogs that frequent the water. We may confidently infer that no man ever selected his water-dogs by the extent to which the skin was developed between their toes; but what he does, is to preserve and breed from those individuals which hunt best in the water, or best retrieve wounded game, and thus he unconsciously selects dogs with feet slightly better webbed. The effects of use from the frequent stretching apart of the toes will likewise aid in the result. Man thus closely imitates Natural Selection. We have an excellent illustration of this same process in North America, where, according to Sir J. Richardson (1/81. 'Fauna Boreali-Americana' 1829 page 62.), all the wolves, foxes, and aboriginal domestic dogs have their feet broader than in the corresponding species of the Old World, and "well calculated for running on the snow." Now, in these Arctic regions, the life or death of every animal will often depend on its success in hunting over the snow when soft; and this will in part depend on the feet being broad; yet they must not be so broad as to interfere with the activity of the animal when the ground is sticky, or with its power of burrowing holes, or with other necessary habits of life.

As changes in domestic breeds which take place so slowly are not to be noticed at any one period, whether due to the selection of individual variations or of differences resulting from crosses, are most important in understanding the origin of our domestic productions, and likewise in throwing indirect light on the changes effected under nature, I will give in detail such cases as I have been able to collect. Lawrence (1/82. 'The Horse in all his Varieties, etc.' 1829 pages 230, 234.), who paid particular attention to the history of the foxhound, writing in 1829, says that between eighty and ninety years before "an entirely new foxhound was raised through the breeder's art," the ears of the old southern hound being reduced, the bone and bulk lightened, the waist increased in length, and the stature somewhat added to. It is believed that this was effected by a cross with a greyhound. With respect to this latter dog, Youatt (1/83. 'The Dog' 1845 pages 31, 35; with respect to King Charles' spaniel page 45; for the setter page 90.), who is generally cautious in his statements, says that the greyhound within the last fifty years, that is before the commencement of the present century, "assumed a somewhat different character from that which he once possessed. He is now distinguished by a beautiful symmetry of form, of which he could not once boast, and he has even superior speed to that which he formerly exhibited. He is no longer used to struggle with deer, but contends with his fellows over a shorter and speedier course." An able writer (1/84. In the 'Encyclop. of Rural Sports' page 557.) believes that our English greyhounds are the descendants, PROGRESSIVELY IMPROVED, of the large rough greyhounds which existed in Scotland so early as the third century. A cross at some former period with the Italian greyhound has been suspected; but this seems hardly probable, considering the feebleness of this latter breed. Lord Orford, as is well-known, crossed his famous greyhounds, which failed in courage, with a bulldog — this breed being chosen from being erroneously supposed to be deficient in the power of scent; "after the sixth or seventh generation," says Youatt, "there was not a vestige left of the form of the bulldog, but his courage and indomitable perseverance remained."

Youatt infers, from a comparison of an old picture of King Charles's spaniels with the living dog, that "the breed of the present day is materially altered for the worse: " the muzzle has become shorter, the forehead more prominent, and the eyes larger; the changes in this case have probably been due to simple selection. The setter, as this author remarks in another place, "is evidently the large spaniel improved to his present peculiar size and beauty, and taught another way of marking his game. If the form of the dog were not sufficiently satisfactory on this point, we might have recourse to history: " he then refers to a document dated 1685 bearing on this subject, and adds that the pure Irish setter shows no signs of a cross with the pointer, which some authors suspect has been the case with the English setter. The bulldog is an English breed, and as I hear from Mr. G.R. Jesse (1/85. Author of 'Researches into the History of the British Dog.), seems to have originated from the mastiff since the time of Shakspeare; but certainly existed in 1631, as shown by Prestwick Eaton's letters. There can be no doubt that the fancy bulldogs of the present day, now that they are not used for bull-baiting, have become greatly reduced in size, without any express intention on the part of the breeder. Our pointers are certainly descended from a Spanish breed, as even their present names, Don, Ponto, Carlos, etc., show; it is said that they were not known in England before the Revolution in 1688 (1/86. See Col. Hamilton Smith on the antiquity of the Pointer, in 'Nat. Lib.' volume 10 page 196.); but the breed since its introduction has been much modified, for Mr. Borrow, who is a sportsman and knows Spain intimately well, informs me that he has not seen in that country any breed "corresponding in figure with the English pointer; but there are genuine pointers near Xeres which have been imported by English gentlemen." A nearly parallel case is offered by the Newfoundland dog, which was certainly brought into England from that country, but which has since been so much modified that, as several writers have observed, it does not now closely resemble any existing native dog in Newfoundland. (1/87. The Newfoundland dog is believed to have originated from a cross between the Esquimaux dog and a large French hound. See Dr. Hodgkin 'British Assoc.' 1844; Bechstein 'Naturgesch. Deutschland' b. 1 s. 574; 'Nat. Lib.' volume 10 page 132; also Mr. Jukes 'Excursion in and about Newfoundland.')

These several cases of slow and gradual changes in our English dogs possess some interest; for though the changes have generally, but not invariably, been caused by one or two crosses with a distinct breed, yet we may feel sure, from the well-known extreme variability of crossed breeds, that rigorous and long-continued selection must have been practised, in order to improve them in a definite manner. As soon as any strain or family became slightly improved or better adapted to alter circumstances, it would tend to supplant the older and less improved strains. For instance, as soon as the old foxhound was improved by a cross with the greyhound, or by simple selection, and assumed its present character — and the change was probably desired owing to the increased fleetness of our hunters — it rapidly spread throughout the country, and is now everywhere nearly uniform. But the process of improvement is still going on for every one tries to improve his strain by occasionally procuring dogs from the best kennels. Through this process of gradual substitution the old English hound has been lost; and so it has been with the Irish wolf-dog, the old English bulldog, and several other breeds, such as the alaunt, as I am informed by Mr. Jesse. But the extinction of former breeds is apparently aided by another cause; for whenever a breed is kept in scanty numbers, as at present with the bloodhound, it is reared with some difficulty, apparently from the evil effects of long-continued close interbreeding. As several breeds of the dog have been slightly but sensibly modified within so short a period as the last one or two centuries, by the selection of the best individuals, modified in many cases by crosses with other breeds; and as we shall hereafter see that the breeding of dogs was attended to in ancient times, as it still is by savages, we may conclude that we have in selection, even if only occasionally practised, a potent means of modification.

DOMESTIC CATS.

Cats have been domesticated in the East from an ancient period; Mr. Blyth informs me that they are mentioned in a Sanskrit writing 2000 years old, and in Egypt their antiquity is known to be even greater, as shown by monumental drawings and their mummied bodies. These mummies, according to De Blainville (1/88. De Blainville 'Osteographie, Felis' page 65 on the character of F. caligulata; pages 85, 89, 90, 175, on the other mummied species. He quotes Ehrenberg on F. maniculata being mummied.), who has particularly studied the subject, belong to no less than three species, namely, F. caligulata, bubastes, and chaus. The two former species are said to be still found, both wild and domesticated, in parts of Egypt. F. caligulata presents a difference in the first inferior milk molar tooth, as compared with the domestic cats of Europe, which makes De Blainville conclude that it is not one of the parent-forms of our cats. Several naturalists, as Pallas, Temminck, Blyth, believe that domestic cats are the descendants of several species commingled: it is certain that cats cross readily with various wild species, and it would appear that the character of the domestic breeds has, at least in some cases, been thus affected. Sir W. Jardine has no doubt that, "in the north of Scotland, there has been occasional crossing with our native species (F. sylvestris), and that the result of these crosses has been kept in our houses. I have seen," he adds, "many cats very closely resembling the wild cat, and one or two that could scarcely be distinguished from it." Mr. Blyth (1/89. Asiatic Soc. of Calcutta; Curator's Report, August 1856. The passage from Sir W. Jardine is quoted from this Report. Mr. Blyth, who has especially attended to the wild and domestic cats of India, has given in this Report a very interesting discussion on their origin.) remarks on this passage, "but such cats are never seen in the southern parts of England; still, as compared with any Indian tame cat, the affinity of the ordinary British cat to F. sylvestris is manifest; and due I suspect to frequent intermixture at a time when the tame cat was first introduced into Britain and continued rare, while the wild species was far more abundant than at present." In Hungary, Jeitteles (1/90. 'Fauna Hungariae Sup.' 1862 s. 12.) was assured on trustworthy authority that a wild male cat crossed with a female domestic cat, and that the hybrids long lived in a domesticated state. In Algiers the domestic cat has crossed with the wild cat (F. lybica) of that country. (1/91. Isid. Geoffroy Saint-Hilaire 'Hist. Nat. Gen.' tome 3 page 177.) In South Africa as Mr. E. Layard informs me, the domestic cat intermingles freely with the wild F. caffra; he has seen a pair of hybrids which were quite tame and particularly attached to the lady who brought them up; and Mr. Fry has found that these hybrids are fertile. In India the domestic cat, according to Mr. Blyth, has crossed with four Indian species. With respect to one of these species, F. chaus, an excellent observer, Sir W. Elliot, informs me that he once killed, near Madras, a wild brood, which were evidently hybrids from the domestic cat; these young animals had a thick lynx-like tail and the broad brown bar on the inside of the forearm characteristic of F. chaus. Sir W. Elliot adds that he has often observed this same mark on the forearms of domestic cats in India. Mr. Blyth states that domestic cats coloured nearly like F. chaus, but not resembling that species in shape, abound in Bengal; he adds, "such a colouration is utterly unknown in European cats, and the proper tabby markings (pale streaks on a black ground, peculiarly and symmetrically disposed), so common in English cats, are never seen in those of India." Dr. D. Short has assured Mr. Blyth (1/92. 'Proc. Zoolog. Soc.' 1863 page 184.) that, at Hansi, hybrids between the common cat and F. ornata (or torquata) occur, "and that many of the domestic cats of that part of India were undistinguishable from the wild F. ornata." Azara states, but only on the authority of the inhabitants, that in Paraguay the cat has crossed with two native species. From these several cases we see that in Europe, Asia, Africa, and America, the common cat, which lives a freer life than most other domesticated animals, has crossed with various wild species; and that in some instances the crossing has been sufficiently frequent to affect the character of the breed.

Whether domestic cats have descended from several distinct species, or have only been modified by occasional crosses, their fertility, as far as is known, is unimpaired. The large Angora or Persian cat is the most distinct in structure and habits of all the domestic breeds; and is believed by Pallas, but on no distinct evidence, to be descended from the F. manul of middle Asia; and I am assured by Mr. Blyth that the Angora cat breeds freely with Indian cats, which, as we have already seen, have apparently been much crossed with F. chaus. In England half-bred Angora cats are perfectly fertile with one another.

Within the same country we do not meet with distinct races of the cat, as we do of dogs and of most other domestic animals; though the cats of the same country present a considerable amount of fluctuating variability. The explanation obviously is that, from their nocturnal and rambling habits, indiscriminate crossing cannot without much trouble be prevented. Selection cannot be brought into play to produce distinct breeds, or to keep those distinct which have been imported from foreign lands. On the other hand, in islands and in countries completely separated from each other, we meet with breeds more or less distinct; and these cases are worth giving, showing that the scarcity of distinct races in the same country is not caused by a deficiency of variability in the animal. The tailless cats of the Isle of Man are said to differ from common cats not only in the want of a tail, but in the greater length of their hind legs, in the size of their heads, and in habits. The Creole cat of Antigua, as I am informed by Mr. Nicholson, is smaller, and has a more elongated head, than the British cat. In Ceylon, as Mr. Thwaites writes to me, every one at first notices the different appearance of the native cat from the English animal; it is of small size, with closely lying hairs; its head is small, with a receding forehead; but the ears are large and sharp; altogether it has what is there called a "low-caste" appearance. Rengger (1/93. 'Saugethiere von Paraguay' 1830 s. 212.) says that the domestic cat, which has been bred for 300 years in Paraguay, presents a striking difference from the European cat; it is smaller by a fourth, has a more lanky body, its hair is short, shining, scanty and lies close, especially on the tail: he adds that the change has been less at Ascension, the capital of Paraguay, owing to the continual crossing with newly imported cats; and this fact well illustrates the importance of separation. The conditions of life in Paraguay appear not to be highly favourable to the cat, for, though they have run half-wild, they do not become thoroughly feral, like so many other European animals. In another part of South America, according to Roulin (1/94. 'Mem. presentes par divers Savans: Acad. Roy. des Sciences' tome 6 page 346. Gomara first noticed this fact in 1554.), the introduced cat has lost the habit of uttering its hideous nocturnal howl. The Rev. W.D. Fox purchased a cat in Portsmouth, which he was told came from the coast of Guinea; its skin was black and wrinkled, fur bluish-grey and short, its ears rather bare, legs long, and whole aspect peculiar. This "negro" cat was fertile with common cats. On the opposite coast of Africa, at Mombas, Captain Owen, R.N. (1/95. 'Narrative of Voyages' volume 2 page 180.) states that all the cats are covered with short stiff hair instead of fur: he gives a curious account of a cat from Algoa Bay, which had been kept for some time on board and could be identified with certainty; this animal was left for only eight weeks at Mombas, but during that short period it "underwent a complete metamorphosis, having parted with its sandy-coloured fur." A cat from the Cape of Good Hope has been described by Desmarest as remarkable from a red stripe extending along the whole length of its back. Throughout an immense area, namely, the Malayan archipelago, Siam, Pegu, and Burmah, all the cats have truncated tails about half the proper length (1/96. J. Crawfurd 'Descript. Dict. of the Indian Islands' page 255. The Madagascar cat is said to have a twisted tail; see Desmarest in 'Encyclop. Nat. Mamm.' 1820 page 233, for some of the other breeds.), often with a sort of knot at the end. In the Caroline archipelago the cats have very long legs, and are of a reddish-yellow colour. (1/97. Admiral Lutke's Voyage volume 3 page 308.) In China a breed has drooping ears. At Tobolsk, according to Gmelin, there is a red-coloured breed. In Asia, also, we find the well-known Angora or Persian breed.

The domestic cat has run wild in several countries, and everywhere assumes, as far as can be judged by the short recorded descriptions, a uniform character. Near Maldonado, in La Plata, I shot one which seemed perfectly wild; it was carefully examined by Mr. Waterhouse (1/98. 'Zoology of the Voyage of the Beagle, Mammalia' page 20. Dieffenbach 'Travels in New Zealand' volume 2 page 185. Ch. St. John 'Wild Sports of the Highlands' 1846 page 40.), who found nothing remarkable in it, excepting its great size. In New Zealand according to Dieffenbach, the feral cats assume a streaky grey colour like that of wild cats; and this is the case with the half-wild cats of the Scotch Highlands.

We have seen that distant countries possess distinct domestic races of the cat. The differences may be in part due to descent from several aboriginal species, or at least to crosses with them. In some cases, as in Paraguay, Mombas, and Antigua, the differences seem due to the direct action of different conditions of life. In other cases some slight effect may possibly be attributed to natural selection, as cats in many cases have largely to support themselves and to escape diverse dangers. But man, owing to the difficulty of pairing cats, has done nothing by methodical selection; and probably very little by unintentional selection; though in each litter he generally saves the prettiest, and values most a good breed of mouse- or rat-catchers. Those cats which have a strong tendency to prowl after game, generally get destroyed by traps. As cats are so much petted, a breed bearing the same relation to other cats, that lapdogs bear to larger dogs, would have been much valued; and if selection could have been applied, we should certainly have had many breeds in each long-civilised country, for there is plenty of variability to work upon.

We see in this country considerable diversity in size, some in the proportions of the body, and extreme variability in colouring. I have only lately attended to this subject, but have already heard of some singular cases of variation; one of a cat born in the West Indies toothless, and remaining so all its life. Mr. Tegetmeier has shown me the skull of a female cat with its canines so much developed that they protruded uncovered beyond the lips; the tooth with the fang being .95, and the part projecting from the gum .6 of an inch in length. I have heard of several families of six-toed cats, in one of which the peculiarity had been transmitted for at least three generations. The tail varies greatly in length; I have seen a cat which always carried its tail flat on its back when pleased. The ears vary in shape, and certain strains, in England, inherit a pencil-like tuft of hairs, above a quarter of an inch in length, on the tips of their ears; and this same peculiarity, according to Mr. Blyth, characterises some cats in India. The great variability in the length of the tail and the lynx-like tufts of hairs on the ears are apparently analogous to differences in certain wild species of the genus. A much more important difference, according to Daubenton (1/99. Quoted by Isid. Geoffroy 'Hist. Nat. Gen.' tome 3 page 427.), is that the intestines of domestic cats are wider, and a third longer, than in wild cats of the same size; and this apparently has been by their less strictly carnivorous diet.

CHAPTER 1.II

HORSES AND ASSES.

HORSE. DIFFERENCES IN THE BREEDS. INDIVIDUAL VARIABILITY OF. DIRECT EFFECTS OF THE CONDITIONS OF LIFE. CAN WITHSTAND MUCH COLD. BREEDS MUCH MODIFIED BY SELECTION. COLOURS OF THE HORSE. DAPPLING. DARK STRIPES ON THE SPINE, LEGS, SHOULDERS, AND FOREHEAD. DUN-COLOURED HORSES MOST FREQUENTLY STRIPED. STRIPES PROBABLY DUE TO REVERSION TO THE PRIMITIVE STATE OF THE HORSE.

ASSES. BREEDS OF. COLOUR OF. LEG- AND SHOULDER-STRIPES. SHOULDER-STRIPES SOMETIMES ABSENT, SOMETIMES FORKED.

The history of the Horse is lost in antiquity. Remains of this animal in a domesticated condition have been found in the Swiss lake-dwellings, belonging to the Neolithic period. (2/1. Rutimeyer 'Fauna der Pfahlbauten' 1861 s. 122.) At the present time the number of breeds is great, as may be seen by consulting any treatise on the Horse. (2/2. See 'Youatt on the Horse': J. Lawrence on the Horse 1829; W.C.L. Martin 'History of the Horse' 1845: Col. H. Smith in 'Nat. Library, Horses' 1841 volume 12: Prof. Veith 'Die naturgesch. Haussaugethiere' 1856.) Looking only to the native ponies of Great Britain, those of the Shetland Isles, Wales, the New Forest, and Devonshire are distinguishable; and so it is, amongst other instances, with each separate island in the great Malay archipelago. (2/3. Crawfurd 'Descript. Dict. of Indian Islands' 1856 page 153. "There are many different breeds, every island having at least one peculiar to it." Thus in Sumatra there are at least two breeds; in Achin and Batubara one; in Java several breeds; one in Bali, Lomboc, Sumbawa (one of the best breeds), Tambora, Bima, Gunung-api, Celebes, Sumba, and Philippines. Other breeds are specified by Zollinger in the 'Journal of the Indian Archipelago' volume 5 page 343 etc.) Some of the breeds present great differences in size, shape of ears, length of mane, proportions of the body, form of the withers and hind quarters, and especially in the head. Compare the race- horse, dray-horse, and a Shetland pony in size, configuration, and disposition; and see how much greater the difference is than between the seven or eight other living species of the genus Equus.

Of individual variations not known to characterise particular breeds, and not great or injurious enough to be called monstrosities, I have not collected many cases. Mr. G. Brown, of the Cirencester Agricultural College, who has particularly attended to the dentition of our domestic animals, writes to me that he has "several times noticed eight permanent incisors instead of six in the jaw." Male horses only should have canines, but they are occasionally found in the mare, though a small size. (2/4. 'The Horse' etc. by John Lawrence 1829 page 14.) The number of ribs on each side is properly eighteen, but Youatt (2/5. 'The Veterinary' London volume 5 page 543.) asserts that not unfrequently there are nineteen, the additional one being always the posterior rib. It is a remarkable fact that the ancient Indian horse is said in the Rig-Veda to have only seventeen ribs; and M. Pietrement (2/6. 'Memoire sur les chevaux a trente-quatre cotes' 1871.), who has called attention to this subject, gives various reasons for placing full trust in this statement, more especially as during former times the Hindoos carefully counted the bones of animals. I have seen several notices of variations in the bones of the leg; thus Mr. Price (2/7. 'Proc. Veterinary Assoc.' in 'The Veterinary' volume 13 page 42.) speaks of an additional bone in the hock, and of certain abnormal appearances between the tibia and astragalus, as quite common in Irish horses, and not due to disease. Horses have often been observed, according to M. Gaudry (2/8. 'Bulletin de la Soc. Geolog.' tome 22 1866 page 22.), to possess a trapezium and a rudiment of a fifth metacarpal bone, so that "one sees appearing by monstrosity, in the foot of the horse, structures which normally exist in the foot of the Hipparion," — an allied and extinct animal. In various countries horn-like projections have been observed on the frontal bones of the horse: in one case described by Mr. Percival they arose about two inches above the orbital processes, and were "very like those in a calf from five to six months old," being from half to three- quarters of an inch in length. (2/9. Mr. Percival of the Enniskillen Dragoons in 'The Veterinary' volume 1 page 224: see Azara, 'Des Quadrupedes du Paraguay' tome 2 page 313. The French translator of Azara refers to other cases mentioned by Huzard as having occurred in Spain.) Azara has described two cases in South America in which the projections were between three and four inches in length: other instances have occurred in Spain.

That there has been much inherited variation in the horse cannot be doubted, when we reflect on the number of the breeds existing throughout the world or even within the same country, and when we know that they have largely increased in number since the earliest known records. (2/10. Godron, 'De l'Espece' tome 1 page 378.) Even in so fleeting a character as colour, Hofacker (2/11. 'Ueber die Eigenschaften' etc. 1828 s. 10.) found that, out of 216 cases in which horses of the same colour were paired, only eleven pairs produced foals of a quite different colour. As Professor Low (2/12. 'Domesticated Animals of the British Islands' pages 527, 532. In all the veterinary treatises and papers which I have read, the writers insist in the strongest terms on the inheritance by the horse of all good and bad tendencies and qualities. Perhaps the principle of inheritance is not really stronger in the horse than in any other animal; but, from its value, the tendency has been more carefully observed.) has remarked, the English race-horse offers the best possible evidence of inheritance. The pedigree of a race-horse is of more value in judging of its probable success than its appearance: "King Herod" gained in prizes 201,505 pounds sterling, and begot 497 winners; "Eclipse" begot 334 winners.

Whether the whole amount of difference between the various breeds has arisen under domestication is doubtful. From the fertility of the most distinct breeds (2/13. Andrew Knight crossed breeds so different in size as a dray-horse and Norwegian pony: see A. Walker on 'Intermarriage' 1838 page 205.) when crossed, naturalists have generally looked at all the breeds as having descended from a single species. Few will agree with Colonel H. Smith, who believes that they have descended from no less than five primitive and differently coloured stocks. (2/14. 'Nat. Library, Horses' volume 12 page 208.) But as several species and varieties of the horse existed (2/15. Gervais 'Hist. Nat. Mamm.' tome 2 page 143. Owen 'British Fossil Mammals' page 383.) during the later tertiary periods, and as Rutimeyer found differences in the size and form of the skull in the earliest known domesticated horses (2/16. 'Kenntniss der fossilen Pferde' 1863 s. 131.), we ought not to feel sure that all our breeds are descended from a single species. The savages of North and South America easily reclaim the feral horses, so that there is no improbability in savages in various quarters of the world having domesticated more than one native species or natural race. M. Sanson (2/17. 'Comptes rendus' 1866 page 485 and 'Journal de l'Anat. et de la Phys.' Mai 1868.) thinks that he has proved that two distinct species have been domesticated, one in the East, and one in North Africa; and that these differed in the number of their lumbar vertebra and in various other parts; but M. Sanson seems to believe that osteological characters are subject to very little variation, which is certainly a mistake. At present no aboriginal or truly wild horse is positively known to exist; for it is commonly believed that the wild horses of the East are escaped domestic animals. (2/18. Mr. W.C.L. Martin, 'The Horse' 1845 page 34, in arguing against the belief that the wild Eastern horses are merely feral, has remarked on the improbability of man in ancient times having extirpated a species in a region where it can now exist in numbers.) If therefore our domestic breeds are descended from several species or natural races, all have become extinct in the wild state.

With respect to the causes of the modifications which horses have undergone, the conditions of life seem to produce a considerable direct effect. Mr. D. Forbes, who has had excellent opportunities of comparing the horses of Spain with those of South America, informs me that the horses of Chile, which have lived under nearly the same conditions as their progenitors in Andalusia, remain unaltered, whilst the Pampas horses and the Puno horses are considerably modified. There can be no doubt that horses become greatly reduced in size and altered in appearance by living on mountains and islands; and this apparently is due to want of nutritious or varied food. Every one knows how small and rugged the ponies are on the Northern islands and on the mountains of Europe. Corsica and Sardinia have their native ponies; and there were (2/19. 'Transact. Maryland Academy' volume 1 part 1 page 28.), or still are, on some islands on the coast of Virginia, ponies like those of the Shetland Islands, which are believed to have originated through exposure to unfavourable conditions. The Puno ponies, which inhabit the lofty regions of the Cordillera, are, as I hear from Mr. D. Forbes, strange little creatures, very unlike their Spanish progenitors. Further south, in the Falkland Islands, the offspring of the horses imported in 1764 have already so much deteriorated in size (2/20. Mr. Mackinnon 'The Falkland Islands' page 25. The average height of the Falkland horses is said to be 14 hands 2 inches. See also my 'Journal of Researches.') and strength that they are unfitted for catching wild cattle with the lasso; so that fresh horses have to be brought for this purpose from La Plata at a great expense. The reduced size of the horses bred on both southern and northern islands, and on several mountain-chains, can hardly have been caused by the cold, as a similar reduction has occurred on the Virginian and Mediterranean islands. The horse can withstand intense cold, for wild troops live on the plains of Siberia under lat. 56 deg, (2/21. Pallas 'Act. Acad. St. Petersburgh' 1777 part 2 page 265. With respect to the tarpans scraping away the snow see Col. Hamilton Smith in 'Nat. Lib.' volume 12 page 165.) and aboriginally the horses must have inhabited countries annually covered with snow, for he long retains the instinct of scraping it away to get at the herbage beneath. The wild tarpans in the East have this instinct; and so it is, as I am informed by Admiral Sulivan, with the horses recently and formerly introduced into the Falkland Islands from La Plata, some of which have run wild; this latter fact is remarkable, as the progenitors of these horses could not have followed this instinct during many generations in La Plata. On the other hand, the wild cattle of the Falklands never scrape away the snow, and perish when the ground is long covered. In the northern parts of America the horses descended from those introduced by the Spanish conquerors of Mexico, have the same habit, as have the native bisons, but not so the cattle introduced from Europe. (2/22. Franklin 'Narrative' volume 1 page 87 note by Sir J. Richardson.)

The horse can flourish under intense heat as well as under intense cold, for he is known to come to the highest perfection, though not attaining a large size, in Arabia and northern Africa. Much humidity is apparently more injurious to the horse than heat or cold. In the Falkland Islands, horses suffer much from the dampness; and this circumstance may perhaps partly account for the singular fact that to the eastward of the Bay of Bengal (2/23. Mr. J.H. Moor 'Notices of the Indian Archipelago' Singapore 1837 page 189. A pony from Java was sent ('Athenaeum' 1842 page 718) to the Queen only 28 inches in height. For the Loo Choo Islands, see Beechey 'Voyage' 4th. edition volume 1 page 499.), over an enormous and humid area, in Ava, Pegu, Siam, the Malayan archipelago, the Loo Choo Islands, and a large part of China, no full-sized horse is found. When we advance as far eastward as Japan, the horse reacquires his full size. (2/24. J. Crawfurd, 'History of the Horse' 'Journal of Royal United Service Institution' volume 4.)

With most of our domesticated animals, some breeds are kept on account of their curiosity or beauty; but the horse is valued almost solely for its utility. Hence semi-monstrous breeds are not preserved; and probably all the existing breeds have been slowly formed either by the direct action of the conditions of life, or through the selection of individual differences. No doubt semi-monstrous breeds might have been formed: thus Mr. Waterton records (2/25. 'Essays on Natural History' 2nd series page 161.) the case of a mare which produced successively three foals without tails; so that a tailless race might have been formed like the tailless races of dogs and cats. A Russian breed of horses is said to have curled hair, and Azara (2/26. 'Quadrupedes du Paraguay' tome 2 page 333. Dr. Canfield informs me that a breed with curly hair was formed by selection at Los Angeles in North America.) relates that in Paraguay horses are occasionally born, but are generally destroyed, with hair like that on the head of a negro; and this peculiarity is transmitted even to half-breeds: it is a curious case of correlation that such horses have short manes and tails, and their hoofs are of a peculiar shape like those of a mule.

It is scarcely possible to doubt that the long-continued selection of qualities serviceable to man has been the chief agent in the formation of the several breeds of the horse. Look at a dray-horse, and see how well adapted he is to draw heavy weights, and how unlike in appearance to any allied wild animal. The English race-horse is known to be derived from the commingled blood of Arabs, Turks, and Barbs; but selection, which was carried on during very early times in England (2/27. See the evidence on this head in 'Land and Water' May 2, 1868.), together with training, have made him a very different animal from his parent-stocks. As a writer in India, who evidently knows the pure Arab well, asks, who now, "looking at our present breed of race-horses, could have conceived that they were the result of the union of the Arab horse and African mare?" The improvement is so marked that in running for the Goodwood Cup "the first descendants of Arabian, Turkish, and Persian horses, are allowed a discount of 18 pounds weight; and when both parents are of these countries a discount of 36 pounds (2/28. Prof. Low 'Domesticated Animals' page 546. With respect to the writer in India see 'India Sporting Review' volume 2 page 181. As Lawrence has remarked ('The Horse' page 9), "perhaps no instance has ever occurred of a three-part bred horse (i.e. a horse, one of whose grandparents was of impure blood) saving his distance in running two miles with thoroughbred racers." Some few instances are on record of seven-eights racers having been successful.) It is notorious that the Arabs have long been as careful about the pedigree of their horses as we are, and this implies great and continued care in breeding. Seeing what has been done in England by careful breeding, can we doubt that the Arabs must likewise have produced during the course of centuries a marked effect on the qualities of their horses? But we may go much farther back in time, for in the Bible we hear of studs carefully kept for breeding, and of horses imported at high prices from various countries. (2/29. Prof. Gervais 'Hist. Nat. Mamm.' tome 2 page 144 has collected many facts on this head. For instance Solomon (1 Kings x. 28) bought horses in Egypt at a high price.) We may therefore conclude that, whether or not the various existing breeds of the horse have proceeded from one or more aboriginal stocks, yet that a great amount of change has resulted from the direct action of the conditions of life, and probably a still greater amount from the long-continued selection by man of slight individual differences.

With several domesticated quadrupeds and birds, certain coloured marks are either strongly inherited or tend to reappear after having been lost for a long time. As this subject will hereafter be seen to be of importance, I will give a full account of the colouring of horses. All English breeds, however unlike in size and appearance, and several of those in India and the Malay archipelago, present a similar range and diversity of colour. The English race-horse, however, is said (2/30. 'The Field' July 13, 1861 page 42.) never to be dun-coloured; but as dun and cream-coloured horses are considered by the Arabs as worthless, "and fit only for Jews to ride" (2/31. E. Vernon Harcourt 'Sporting in Algeria' page 26.), these tints may have been removed by long-continued selection. Horses of every colour, and of such widely different kinds as dray-horses, cobs, and ponies, are all occasionally dappled (2/32. I state this from my own observations made during several years on the colours of horses. I have seen cream-coloured, light-dun and mouse-dun horses dappled, which I mention because it has been stated (Martin 'History of the Horse' page 134) that duns are never dappled. Martin (page 205) refers to dappled asses. In the 'Farrier' (London 1828 pages 453, 455) there are some good remarks on the dappling of horses; and likewise in Col. Hamilton Smith on 'The Horse.'), in the same manner as is so conspicuous with grey horses. This fact does not throw any clear light on the colouring of the aboriginal horse, but is a case of analogous variation, for even asses are sometimes dappled, and I have seen, in the British Museum, a hybrid from the ass and zebra dappled on its hinder quarters. By the expression analogous variation (and it is one that I shall often have occasion to use) I mean a variation occurring in a species or variety which resembles a normal character in another and distinct species or variety. Analogous variations may arise, as will be explained in a future chapter, from two or more forms with a similar constitution having been exposed to similar conditions, — or from one of two forms having reacquired through reversion a character inherited by the other form from their common progenitor, — or from both forms having reverted to the same ancestral character. We shall immediately see that horses occasionally exhibit a tendency to become striped over a large part of their bodies; and as we know that in the varieties of the domestic cat and in several feline species stripes readily pass into spots and cloudy marks — even the cubs of the uniformly-coloured lion being spotted with dark marks on a lighter ground — we may suspect that the dappling of the horse, which has been noticed by some authors with surprise, is a modification or vestige of a tendency to become striped.

(FIGURE 1. DUN DEVONSHIRE PONY, with shoulder, spinal, and leg stripes.)

[This tendency in the horse to become striped is in several respects an interesting fact. Horses of all colours, of the most diverse breeds, in various parts of the world, often have a dark stripe extending along the spine, from the mane to the tail; but this is so common that I need enter into no particulars. (2/33. Some details are given in 'The Farrier' 1828 pages 452, 455. One of the smallest ponies I ever saw, of the colour of a mouse, had a conspicuous spinal stripe. A small Indian chestnut pony had the same stripe, as had a remarkably heavy chestnut cart-horse. Race-horses often have the spinal stripe.) Occasionally horses are transversely barred on the legs, chiefly on the under side; and more rarely they have a distinct stripe on the shoulder, like that on the shoulder of the ass, or a broad dark patch representing a stripe. Before entering on any details I must premise that the term dun-coloured is vague, and includes three groups of colours, viz., that between cream-colour and reddish-brown, which graduates into light-bay or light-chestnut — this, I believe is often called fallow-dun; secondly, leaden or slate-colour or mouse-dun, which graduates into an ash-colour; and, lastly, dark-dun, between brown and black. In England I have examined a rather large, lightly-built, fallow-dun Devonshire pony (Figure 1), with a conspicuous stripe along the back, with light transverse stripes on the under sides of its front legs, and with four parallel stripes on each shoulder. Of these four stripes the posterior one was very minute and faint; the anterior one, on the other hand, was long and broad, but interrupted in the middle, and truncated at its lower extremity, with the anterior angle produced into a long tapering point. I mention this latter fact because the shoulder-stripe of the ass occasionally presents exactly the same appearance. I have had an outline and description sent to me of a small, purely-bred, light fallow-dun Welch pony, with a spinal stripe, a single transverse stripe on each leg, and three shoulder-stripes; the posterior stripe corresponding with that on the shoulder of the ass was the longest, whilst the two anterior parallel stripes, arising from the mane, decreased in length, in a reversed manner as compared with the shoulder-stripes on the above-described Devonshire pony. I have seen a bright fallow-dun cob, with its front legs transversely barred on the under sides in the most conspicuous manner; also a dark- leaden mouse-coloured pony with similar leg stripes, but much less conspicuous; also a bright fallow-dun colt, fully three-parts thoroughbred, with very plain transverse stripes on the legs; also a chestnut-dun cart- horse with a conspicuous spinal stripe, with distinct traces of shoulder- stripes, but none on the legs; I could add other cases. My son made a sketch for me of a large, heavy, Belgian cart-horse, of a fallow-dun, with a conspicuous spinal stripe, traces of leg-stripes, and with two parallel (three inches apart) stripes about seven or eight inches in length on both shoulders. I have seen another rather light cart-horse, of a dirty dark cream-colour, with striped legs, and on one shoulder a large ill-defined dark cloudy patch, and on the opposite shoulder two parallel faint stripes. All the cases yet mentioned are duns of various tints; but Mr. W.W. Edwards has seen a nearly thoroughbred chestnut horse which had the spinal stripe, and distinct bars on the legs; and I have seen two bay carriage-horses with black spinal stripes; one of these horses had on each shoulder a light shoulder-stripe, and the other had a broad back ill-defined stripe, running obliquely half-way down each shoulder; neither had leg-stripes.

The most interesting case which I have met with occurred in a colt of my own breeding. A bay mare (descended from a dark-brown Flemish mare by a light grey Turcoman horse) was put to Hercules, a thoroughbred dark bay, whose sire (Kingston) and dam were both bays. The colt ultimately turned out brown; but when only a fortnight old it was a dirty bay, shaded with mouse-grey, and in parts with a yellowish tint: it had only a trace of the spinal stripe, with a few obscure transverse bars on the legs; but almost the whole body was marked with very narrow dark stripes, in most parts so obscure as to be visible only in certain lights, like the stripes which may be seen on black kittens. These stripes were distinct on the hind-quarters, where they diverged from the spine, and pointed a little forwards; many of them as they diverged became a little branched, exactly in the same manner as in some zebrine species. The stripes were plainest on the forehead between the ears, where they formed a set of pointed arches, one under the other, decreasing in size downwards towards the muzzle; exactly similar marks may be seen on the forehead of the quagga and Burchell's zebra. When this foal was two or three months old all the stripes entirely disappeared. I have seen similar marks on the forehead of a fully grown, fallow-dun, cob-like horse, having a conspicuous spinal stripe, and with its front legs well barred.

In Norway the colour of the native horse or pony is dun, varying from almost cream-colour to dark-mouse dun; and an animal is not considered purely bred unless it has the spinal and leg-stripes. (2/34. I have received information, through the kindness of the Consul-General, Mr. J.R. Crowe, from Prof. Boeck, Rasck, and Esmarck, on the colours of the Norwegian ponies. See also 'The Field' 1861 page 431.) My son estimated that about a third of the ponies which he saw there had striped legs; he counted seven stripes on the fore-legs and two on the hind-legs of one pony; only a few of them exhibited traces of shoulder stripes; but I have heard of a cob imported from Norway which had the shoulder as well as the other stripes well developed. Colonel H. Smith (2/35. Col. Hamilton Smith 'Nat. Lib.' volume 12 page 275.) alludes to dun-horses with the spinal stripe in the Sierras of Spain; and the horses originally derived from Spain, in some parts of South America, are now duns. Sir W. Elliot informs me that he inspected a herd of 300 South American horses imported into Madras, and many of these had transverse stripes on the legs and short shoulder-stripes; the most strongly marked individual, of which a coloured drawing was sent me, was a mouse-dun, with the shoulder-stripes slightly forked.

In the North-Western parts of India striped horses of more than one breed are apparently commoner than in any other part of the world; and I have received information respecting them from several officers, especially from Colonel Poole, Colonel Curtis, Major Campbell, Brigadier St. John, and others. The Kattywar horses are often fifteen or sixteen hands in height, and are well but lightly built. They are of all colours, but the several kinds of duns prevail; and these are so generally striped, that a horse without stripes is not considered pure. Colonel Poole believes that all the duns have the spinal stripe, the leg-stripes are generally present, and he thinks that about half the horses have the shoulder-stripe; this stripe is sometimes double or treble on both shoulders. Colonel Poole has often seen stripes on the cheeks and sides of the nose. He has seen stripes on the grey and bay Kattywars when first foaled, but they soon faded away. I have received other accounts of cream-coloured, bay, brown, and grey Kattywar horses being striped. Eastward of India, the Shan (north of Burmah) ponies, as I am informed by Mr. Blyth, have spinal, leg, and shoulder stripes. Sir W. Elliot informs me that he saw two bay Pegu ponies with leg-stripes. Burmese and Javanese ponies are frequently dun-coloured, and have the three kinds of stripes, "in the same degree as in England." (2/36. Mr. G. Clark in 'Annal and Mag. of Nat. History' 2nd series volume 2 1848 page 363. Mr. Wallace informs me that he saw in Java a dun and clay-coloured horse with spinal and leg stripes.) Mr. Swinhoe informs me that he examined two light- dun ponies of two Chinese breeds, viz., those of Shanghai and Amoy; both had the spinal stripe, and the latter an indistinct shoulder-stripe.

We thus see that in all parts of the world breeds of the horse as different as possible, when of a dun-colour (including under this term a wide range of tint from cream to dusty black), and rarely when almost white tinged with yellow, grey, bay, and chestnut, have the several above-specified stripes. Horses which are of a yellow colour with white mane and tail, and which are sometimes called duns, I have never seen with stripes. (2/37. See also on this point 'The Field' July 27, 1861 page 91.)

From reasons which will be apparent in the chapter on Reversion, I have endeavoured, but with poor success, to discover whether duns, which are so much oftener striped than other coloured horses, are ever produced from the crossing of two horses, neither of which are duns. Most persons to whom I have applied believe that one parent must be dun; and it is generally asserted that, when this is the case, the dun-colour and the stripes are strongly inherited. (2/38. 'The Field' 1861 pages 431, 493, 545.) One case, however, has fallen under my own observation of a foal from a black mare by a bay horse, which when fully grown was a dark fallow-dun and had a narrow but plain spinal stripe. Hofacker (2/39. 'Ueber die Eigenschaften' etc. 1828 s. 13, 14.) gives two instances of mouse-duns (Mausrapp) being produced from two parents of different colours and neither duns.

The stripes of all kinds are generally plainer in the foal than in the adult horse, being commonly lost at the first shedding of the hair. (2/40. Von Nathusius 'Vortrage uber Viehzucht' 1872 135.) Colonel Poole believes that "the stripes in the Kattywar breed are plainest when the colt is first foaled; they then become less and less distinct till after the first coat is shed, when they come out as strongly as before; but certainly often fade away as the age of the horse increases." Two other accounts confirm this fading of the stripes in old horses in India. One writer, on the other hand, states that colts are often born without stripes, but that they appear as the colt grows older. Three authorities affirm that in Norway the stripes are less plain in the foal than in the adult. In the case described by me of the young foal which was narrowly striped over nearly all its body, there was no doubt about the early and complete disappearance of the stripes. Mr. W.W. Edwards examined for me twenty-two foals of race-horses, and twelve had the spinal stripe more or less plain; this fact, and some other accounts which I have received, lead me to believe that the spinal stripe often disappears in the English race-horse when old. With natural species, the young often exhibit characters which disappear at maturity.]

The stripes are variable in colour, but are always darker than the rest of the body. They do not by any means always coexist on the different parts of the body: the legs may be striped without any shoulder-stripe, or the converse case, which is rarer, may occur; but I have never heard of either shoulder or leg-stripes without the spinal stripe. The latter is by far the commonest of all the stripes, as might have been expected, as it characterises the other seven or eight species of the genus. It is remarkable that so trifling a character as the shoulder-stripe being double or triple should occur in such different breeds as Welch and Devonshire ponies, the Shan pony, heavy cart-horses, light South American horses, and the lanky Kattywar breed. Colonel Hamilton Smith believes that one of his five supposed primitive stocks was dun-coloured and striped; and that the stripes in all the other breeds result from ancient crosses with this one primitive dun; but it is extremely improbable that different breeds living in such distant quarters of the world should all have been crossed with any one aboriginally distinct stock. Nor have we any reason to believe that the effects of a cross at a very remote period would be propagated for so many generations as is implied on this view.

With respect to the primitive colour of the horse having been dun, Colonel Hamilton Smith (2/41. 'Nat. Library' volume 12 1841 pages 109, 156 to 163, 280, 281. Cream-colour, passing into Isabella (i.e. the colour of the dirty linen of Queen Isabella), seems to have been common in ancient times. See also Pallas's account of the wild horses of the East, who speaks of dun and brown as the prevalent colours. In the Icelandic sagas, which were committed to writing in the twelfth century, dun-coloured horses with a black spinal stripe are mentioned; see Dasent's translation volume 1 page 169.) has collected a large body of evidence showing that this tint was common in the East as far back as the time of Alexander, and that the wild horses of Western Asia and Eastern Europe now are, or recently were, of various shades of dun. It seems that not very long ago a wild breed of dun- coloured horses with a spinal stripe was preserved in the royal parks in Prussia. I hear from Hungary that the inhabitants of that country look at the duns with a spinal stripe as the aboriginal stock, and so it is in Norway. Dun-coloured ponies are not rare in the mountainous parts of Devonshire, Wales, and Scotland, where the aboriginal breed would have the best chance of being preserved. In South America in the time of Azara, when the horse had been feral for about 250 years, 90 out of 100 horses were "bai-chatains," and the remaining ten were "zains," that is brown; not more than one in 2000 being black. In North America the feral horses show a strong tendency to become roans of various shades; but in certain parts, as I hear from Dr. Canfield, they are mostly duns and striped. (2/42. Azara 'Quadrupedes du Paraguay' tome 2 page 307. In North America Catlin (volume 2 page 57) describes the wild horses, believed to have descended from the Spanish horses of Mexico, as of all colours, black, grey, roan, and roan pied with sorrel. F. Michaux 'Travels in North America' English translation page 235, describes two wild horses from Mexico as roan. In the Falkland Islands, where the horse has been feral only between 60 and 70 years, I was told that roans and iron-greys were the prevalent colours. These several facts show that horses do not soon revert to any uniform colour.)

In the following chapters on the Pigeon we shall see that a blue bird is occasionally produced by pure breeds of various colours and that when this occurs certain black marks invariably appear on the wings and tail; so again, when variously coloured breeds are crossed, blue birds with the same black marks are frequently produced. We shall further see that these facts are explained by, and afford strong evidence in favour of, the view that all the breeds are descended from the rock-pigeon, or Columba livia, which is thus coloured and marked. But the appearance of the stripes on the various breeds of the horse, when of a dun colour, does not afford nearly such good evidence of their descent from a single primitive stock as in the case of the pigeon: because no horse certainly wild is known as a standard of comparison; because the stripes when they appear are variable in character; because there is far from sufficient evidence that the crossing of distinct breeds produces stripes, and lastly, because all the species of the genus Equus have the spinal stripe, and several species have shoulder and leg stripes. Nevertheless the similarity in the most distinct breeds in their general range of colour, in their dappling, and in the occasional appearance, especially in duns, of leg-stripes and of double or triple shoulder-stripes, taken together, indicate the probability of the descent of all the existing races from a single, dun-coloured, more or less striped, primitive stock, to which our horses occasionally revert.

THE ASS.

Four species of Asses, besides three zebras, have been described by naturalists. There is now little doubt that our domesticated animal is descended from the Equus taeniopus of Abyssinia. (2/43. Dr. Sclater in 'Proc. Zoolog. Soc.' 1862 page 164. Dr. Hartmann says ('Annalen der Landw.' b. 44 page 222) that this animal in its wild state is not always striped across the legs.) The ass is sometimes advanced as an instance of an animal domesticated, as we know by the Old Testament, from an ancient period, which has varied only in a very slight degree. But this is by no means strictly true; for in Syria alone there are four breeds (2/44. W.C. Martin 'History of the Horse' 1845 page 207.); first, a light and graceful animal, with an agreeable gait, used by ladies; secondly, an Arab breed reserved exclusively for the saddle; thirdly, a stouter animal used for ploughing and various purposes; and lastly, the large Damascus breed, with a peculiarly long body and ears. In the South of France also there are several breeds, and one of extraordinary size, some individuals being as tall as full-sized horses. Although the ass in England is by no means uniform in appearance, distinct breeds have not been formed. This may probably be accounted for by the animal being kept chiefly by poor persons, who do not rear large numbers, nor carefully match and select the young. For, as we shall see in a future chapter, the ass can with ease be greatly improved in size and strength by careful selection, combined no doubt with good food; and we may infer that all its other characters would be equally amenable to selection. The small size of the ass in England and Northern Europe is apparently due far more to want of care in breeding than to cold; for in Western India, where the ass is used as a beast of burden by some of the lower castes, it is not much larger than a Newfoundland dog, "being generally not more than from twenty to thirty inches high." (2/45. Col. Sykes Cat. of Mammalia 'Proc. Zoolog. Soc.' July 12, 1831. Williamson 'Oriental Field Sports' volume 2 quoted by Martin page 206.)

The ass varies greatly in colour; and its legs, especially the fore-legs, both in England and other countries — for instance, in China — are occasionally barred more plainly than those of dun-coloured horses. Thirteen or fourteen transverse stripes have been counted on both the fore and hind legs. With the horse the occasional appearance of leg-stripes was accounted for by reversion to a supposed parent-form, and in the case of the ass we may confidently believe in this explanation, as E. taeniopus is known to be barred, though only in a slight degree, and not quite invariably. The stripes are believed to occur most frequently and to be plainest on the legs of the domestic ass during early youth (2/46. Blyth in 'Charlesworth's Mag. of Nat. Hist.' vol 4 1840 page 83. I have also been assured by a breeder that this is the case.), as likewise occurs with the horse. The shoulder-stripe, which is so eminently characteristic of the species, is nevertheless variable in breadth, length, and manner of termination. I have measured one four times as broad as another, and some more than twice as long as others. In one light-grey ass the shoulder- stripe was only six inches in length, and as thin as a piece of string; and in another animal of the same colour there was only a dusky shade representing a stripe. I have heard of three white asses, not albinoes, with no trace of shoulder or spinal stripes (2/47. One case is given by Martin 'The Horse' page 205.); and I have seen nine other asses with no shoulder-stripe, and some of them had no spinal stripe. Three of the nine were light-greys, one a dark-grey, another grey passing into reddish-roan, and the others were brown, two being tinted on parts of their bodies with a reddish or bay shade. If therefore grey and reddish-brown asses had been steadily selected and bred from, the shoulder stripe would probably have been lost almost as generally and completely as in the case of the horse.

The shoulder stripe on the ass is sometimes double, and Mr. Blyth has seen even three or four parallel stripes. (2/48. 'Journal As. Soc. of Bengal' volume 28 1860 page 231. Martin on the Horse page 205.) I have observed in ten cases shoulder-stripes abruptly truncated at the lower end, with the anterior angle produced into a tapering point, precisely as in the above dun Devonshire pony. I have seen three cases of the terminal portion abruptly and angularly bent; and have seen and heard of four cases of a distinct though slight forking of the stripe. In Syria, Dr. Hooker and his party observed for me no less than five similar instances of the shoulder- stripe plainly bifurcating over the fore leg. In the common mule it likewise sometimes bifurcates. When I first noticed the forking and angular bending of the shoulder-stripe, I had seen enough of the stripes in the various equine species to feel convinced that even a character so unimportant as this had a distinct meaning, and was thus led to attend to the subject. I now find that in the E. burchellii and quagga, the stripe which corresponds with the shoulder-stripe of the ass, as well as some of the stripes on the neck, bifurcate, and that some of those near the shoulder have their extremities bent angularly backwards. The bifurcation and angular bending of the stripes on the shoulders apparently are connected with the nearly upright stripes on the sides of the body and neck changing their direction and becoming transverse on the legs. Finally, we see that the presence of shoulder, leg, and spinal stripes in the horse, — their occasional absence in the ass, — the occurrence of double and triple shoulder-stripes in both animals, and the similar manner in which these stripes terminate downwards, — are all cases of analogous variation in the horse and ass. These cases are probably not due to similar conditions acting on similar constitutions, but to a partial reversion in colour to the common progenitor of the genus. We shall hereafter return to this subject, and discuss it more fully.

CHAPTER 1.III