The Variation of Animals and Plants by Charles Darwin - HTML preview

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Chapter III: Pigs - Cattle - Sheep – Goats

PIGS BELONG TO TWO DISTINCT TYPES, SUS SCROFA AND INDICUS — TORFSCHWEIN — JAPAN PIGS — FERTILITY OF CROSSED PIGS — CHANGES IN THE SKULL OF THE HIGHLY CULTIVATED RACES — CONVERGENCE OF CHARACTER — GESTATION — SOLID-HOOFED SWINE — CURIOUS APPENDAGES TO THE JAWS — DECREASE IN SIZE OF THE TUSKS — YOUNG PIGS LONGITUDINALLY STRIPED — FERAL PIGS — CROSSED BREEDS.

CATTLE — ZEBU A DISTINCT SPECIES — EUROPEAN CATTLE PROBABLY DESCENDED FROM THREE WILD FORMS — ALL THE RACES NOW FERTILE TOGETHER — BRITISH PARK CATTLE — ON THE COLOUR OF THE ABORIGINAL SPECIES — CONSTITUTIONAL DIFFERENCES — SOUTH AFRICAN RACES — SOUTH AMERICAN RACES — NIATA CATTLE — ORIGIN OF THE VARIOUS RACES OF CATTLE.

SHEEP — REMARKABLE RACES OF — VARIATIONS ATTACHED TO THE MALE SEX — ADAPTATIONS TO VARIOUS CONDITIONS — GESTATION OF — CHANGES IN THE WOOL — SEMI-MONSTROUS BREEDS.

GOATS — REMARKABLE VARIATIONS OF.

The breeds of the pig have recently been more closely studied, though much still remains to be done, than those of almost any other domesticated animal. This has been effected by Hermann von Nathusius in two admirable works, especially in the later one on the Skulls of the several races, and by Rütimeyer in his celebrated Fauna of the ancient Swiss lakedwellings. Nathusius has shown that all the known breeds may be divided into two great groups: one resembling in all important respects and no doubt descended from the common wild boar; so that this may be called the Sus scrofa group. The other group differs in several important and constant osteological characters; its wild parent-form is unknown; the name given to it by Nathusius, according to the law of priority, is Sus indicus, of Pallas. This name must now be followed, though an unfortunate one, as the wild aboriginal does not inhabit India, and the best-known domesticated breeds have been imported from Siam and China.

First for the Sus scrofa breeds, or those resembling the common wild boar. These still exist, according to Nathusius ('Schweineschädel' s. 75), in various parts of central and northern Europe; formerly every kingdom, and almost every province in Britain, possessed its own native breed; but these are now everywhere rapidly disappearing, being replaced by improved breeds crossed with the S. indicus form. The skull in the breeds of the S. scrofa type resembles, in all important respects, that of the European wild boar; but it has become ('Schweineschädel' s. 63-68) higher and broader relatively to its length; and the hinder part is more upright. The differences, however, are all variable in degree. The breeds which thus resemble S. scrofa in their essential skull characters differ conspicuously from each other in other respects, as in the length of the ears and legs, curvature of the ribs, colour, hairiness, size and proportions of the body.
The wild Sus scrofa has a wide range, namely, Europe, North Africa, as identified by osteological characters by Rütimeyer, and Hindostan, as similarly identified by Nathusius. But the wild boars inhabiting these several countries differ so much from each other in external characters, that they have been ranked by some naturalists as specifically distinct. Even within Hindostan these animals, according to Mr. Blyth, form very distinct races in the different districts; in the N. Western provinces, as I am informed by the Rev. R. Everest, the boar never exceeds 36 inches in height, whilst in Bengal one has been measured 44 inches in height. In Europe, Northern Africa, and Hindostan, domestic pigs have been known to cross with the wild native species; and in Hindostan an accurate observer,Sir Walter Elliot, after describing the differences between wild Indian and wild German boars, remarks that "the same differences are perceptible in the domesticated individuals of the two countries." We may therefore conclude that the breeds of the Sus scrofa type are descended from, or have been modified by crossing with, forms which may be ranked as geographical races, but which, according to some naturalists, ought to be ranked as distinct species.

Pigs of the Sus indicus type are best known to Englishmen under the form of the Chinese breed. The skull of S. indicus, as described by Nathusius, differs from that of S. scrofa in several minor respects, as in its greater breadth and in some details in the teeth; but chiefly in the shortness of the lachrymal bones, in the greater width of the fore part of the palate-bones, and in the divergence of the premolar teeth. It deserves especial notice that these latter characters are not gained, even in the least degree, by the domesticated forms of S. scrofa. After reading the remarks and descriptions given by Nathusius, it seems to me to be merely playing with words to doubt whether S. indicus ought to be ranked as a species; for the above-specified differences are more strongly marked than any that can be pointed out between, for instance, the fox and the wolf, or the ass and the horse. As already stated, S. indicus is not known in a wild state; but its domesticated forms, according to Nathusius, come near to S. vittatus of Java and some allied species. A pig found wild in the Aru islands ('Schweineschädel' s. 169) is apparently identical with S. indicus; but it is doubtful whether this is a truly native animal. The domesticated breeds of China, Cochin-China, and Siam belong to this type. The Roman or Neapolitan breed, the Andalusian, the Hungarian, and the "Krause" swine of Nathusius, inhabiting southeastern Europe and Turkey, and having fine curly hair, and the small Swiss "Bündtnerschwein" of Rütimeyer, all agree in their more important skull-characters with S. indicus, and, as is supposed, have all been largely crossed with this form. Pigs of this type have existed during a long period on the shores of the Mediterranean, for a figure ('Schweineschädel' s. 142) closely resembling the existing Neapolitan pig was found in the buried city of Herculaneum.

Rütimeyer has made the remarkable discovery that there lived contemporaneously in Switzerland, during the Neolithic period, two domesticated forms, the S. scrofa, and the S. scrofa palustris or Torfschwein. Rütimeyer perceived that the latter approached the Eastern breeds, and, according to Nathusius, it certainly belongs to the S. indicus group; but Rütimeyer has subsequently shown that it differs in some well-marked characters. This author was formerly convinced that his Torfschwein existed as a wild animal during the first part of the Stone period, and was domesticated during a later part of the same period. Nathusius, whilst he fully admits the curious fact first observed by Rütimeyer, that the bones of domesticated and wild animals can be distinguished by their different aspect, yet, from special difficulties in the case of the bones of the pig ('Schweineschädel' s. 147), is not convinced of the truth of the above conclusion; and Rütimeyer himself seems now to feel some doubt. Other naturalists have also argued strongly on the same side as Nathusius.

Several breeds, differing in the proportions of the body, in the length of the ears, in the nature of the hair, in colour, etc., come under the S. indicus type. Nor is this surprising, considering how ancient the domestication of this form has been both in Europe and in China. In this latter country the date is believed by an eminent Chinese scholar to go back at least 4900 years from the present time. This same scholar alludes to the existence of many local varieties of the pig in China; and at the present time the Chinese take extraordinary pains in feeding and tending their pigs, not even allowing them to walk from place to place. Hence these pigs, as Nathusius has remarked, display in an eminent degree the characters of a highly-cultivated race, and hence, no doubt, their high value in the improvement of our European breeds. Nathusius makes a remarkable statement ('Schweineschädel' s. 138), that the infusion of the 1/32nd, or even of the 1/64th, part of the blood of S. indicus into a breed of S. scrofa, is sufficient plainly to modify the skull of the latter species. This singular fact may perhaps be accounted for by several of the chief distinctive characters of S. indicus, such as the shortness of the lachrymal bones, etc., being common to several species of the genus; for in crosses characters which are common to many species apparently tend to be prepotent over those appertaining to only a few species.

The Japan pig ( S. pliciceps of Gray), which was formerly exhibited in the Zoological Gardens, has an extraordinary appearance from its short head, broad forehead and nose, great fleshy ears, and deeply furrowed skin. Figure 2 is copied from that given by Mr. Bartlett. Not only is the face furrowed, but thick folds of skin, which are harder than the other parts, almost like the plates on the Indian rhinoceros, hang about the shoulders and rump. It is coloured black, with white feet, and breeds true. That it has long been domesticated there can be little doubt; and this might have been inferred even from the fact that its young are not longitudinally striped; for this is a character common to all the species included within the genus Sus and the allied genera whilst in their natural state. Dr. Gray has described the skull of this animal, which he ranks not only as a distinct species, but places it in a distinct section of the genus. Nathusius, however, after his careful study of the whole group, states positively ('Schweineschädel' s. 153-158). that the skull in all essential characters closely resembles that of the short-eared Chinese breed of the S. indicus type. Hence Nathusius considers the Japan pig as only a domesticated variety of S. indicus: if this really be the case, it is a wonderful instance of the amount of modification which can be effected under domestication.

Formerly there existed in the central islands of the Pacific Ocean a singular breed of pigs. These are described by the Rev. D. Tyerman and G. Bennett as of small size, humpbacked, with a disproportionately long head, with short ears turned backwards, with a bushy tail not more than two inches in length, placed as if it grew from the back. Within half a century after the introduction of European and Chinese pigs into these islands, the native breed, according to the above authors, became almost completely lost by being repeatedly crossed with them. Secluded islands, as might have been expected, seem favourable for the production or retention of peculiar breeds; thus, in the Orkney Islands, the hogs have been described as very small, with erect and sharp ears, and "with an appearance altogether different from the hogs brought from the south."

Seeing how different the Chinese pigs, belonging to the Sus indicus type, are in their osteological characters and in external appearance from the pigs of the S. scrofa type, so that they must be considered specifically distinct, it is a fact well deserving attention, that Chinese and common pigs have been repeatedly crossed in various manners, with unimpaired fertility. One great breeder who had used pure Chinese pigs assured me that the fertility of the half-breeds inter se and of their recrossed progeny was actually increased; and this is the general belief of agriculturists. Again, the Japan pig or S. pliciceps of Gray is so distinct in appearance from all common pigs, that it stretches one's belief to the utmost to admit that it is simply a domestic variety; yet this breed has been found perfectly fertile with the Berkshire breed; and Mr. Eyton informs me that he paired a half-bred brother and sister and found them quite fertile together.

The modification of the skull in the most highly cultivated races is wonderful. To appreciate the amount of change, Nathusius' work, with its excellent figures, should be studied. The whole of the exterior in all its parts has been altered: the hinder surface, instead of sloping backwards, is directed forwards, entailing many changes in other parts; the front of the head is deeply concave; the orbits have a different shape; the auditory meatus has a different direction and shape; the incisors of the upper and lower jaws do not touch each other, and they stand in both jaws beyond the plane of the molars; the canines of the upper jaw stand in front of those of the lower jaw, and this is a remarkable anomaly: the articular surfaces of the occipital condyles are so greatly changed in shape, that, as Nathusius remarks (s. 133), no naturalist, seeing this important part of the skull by itself, would suppose that it belonged to the genus Sus. These and various other modifications, as Nathusius observes, can hardly be considered as monstrosities, for they are not injurious, and are strictly inherited. The whole head is much shortened; thus, whilst in common breeds its length to that of the body is as 1 to 6, in the "cultur-racen" the proportion is as 1 to 9, and even recently as 1 to 11. The woodcut (Figure 3) of the head of a wild boar and of a sow from a photograph of the Yorkshire Large Breed, may aid in showing how greatly the head in a highly cultivated race has been modified and shortened.

Nathusius has well discussed the causes of the remarkable changes in the skull and shape of the body which the highly cultivated races have undergone. These modifications occur chiefly in the pure and crossed races of the S. indicus type; but their commencement may be clearly detected in the slightly improved breeds of the S. scrofa type.17 Nathusius states positively (s. 99, 103), as the result of common experience and of his experiments, that rich and abundant food, given during youth, tends by some direct action to make the head broader and shorter; and that poor food works a contrary result. He lays much stress on the fact that all wild and semi-domesticated pigs, in ploughing up the ground with their muzzles, have, whilst young, to exert the powerful muscles fixed to the hinder part of the head. In highly cultivated races this habit is no longer followed, and consequently the back of the skull becomes modified in shape, entailing other changes in other parts. There can hardly be a doubt that so great a change in habits would affect the skull; but it seems rather doubtful how far this will account for the greatly reduced length of the skull and for its concave front. It is well known (Nathusius himself advancing many cases, s. 104) that there is a strong tendency in many domestic animals—in bull- and pug-dogs, in the niata cattle, in sheep, in Polish fowls, short-faced tumbler pigeons, and in one variety of the carp—for the bones of the face to become greatly shortened. In the case of the dog, as H. Müller has shown, this seems caused by an abnormal state of the primordial cartilage. We may, however, readily admit that abundant and rich food supplied during many generations would give an inherited tendency to increased size of body, and that, from disuse, the limbs would become finer and shorter. We shall in a future chapter see also that the skull and limbs are apparently in some manner correlated, so that any change in the one tends to affect the other.

Nathusius has remarked, and the observation is an interesting one, that the peculiar form of the skull and body in the most highly cultivated races is not characteristic of any one race, but is common to all when improved up to the same standard. Thus the largebodied, long-eared, English breeds with a convex back, and the small-bodied, shorteared, Chinese breeds with a concave back, when bred to the same state of perfection, nearly resemble each other in the form of the head and body. This result, it appears, is partly due to similar causes of change acting on the several races, and partly to man breeding the pig for one sole purpose, namely, for the greatest amount of flesh and fat; so that selection has always tended towards one and the same end. With most domestic animals the result of selection has been divergence of character, here it has been convergence.

The nature of the food supplied during many generations has apparently affected the length of the intestines; for, according to Cuvier, their length to that of the body in the wild boar is as 9 to 1,—in the common domestic boar as 13·5 to 1,—and in the Siam breed as 16 to 1. In this latter breed the greater length may be due either to descent from a distinct species or to more ancient domestication. The number of mammæ vary, as does the period of gestation. The latest authority says that "the period averages from 17 to 20 weeks," but I think there must be some error in this statement: in M. Tessier's observations on 25 sows it varied from 109 to 123 days. The Rev. W. D. Fox has given me ten carefully recorded cases with well-bred pigs, in which the period varied from 101 to 116 days. According to Nathusius the period is shortest in the races which come early to maturity; but the course of their development does not appear to be actually shortened, for the young animal is born, judging from the state of the skull, less fully developed, or in a more embryonic condition, than in the case of common swine. In the highly cultivated and early matured races the teeth, also, are developed earlier.

The difference in the number of the vertebræ and ribs in different kinds of pigs, as observed by Mr. Eyton, and as given in the following table, has often been quoted. The African sow probably belongs to the S. scrofa type; and Mr. Eyton informs me that, since the publication of this paper, cross-bred animals from the African and English races were found by Lord Hill to be perfectly fertile.

English
Long-legged Male.
African Chinese Wild Boar Female. Male. from Cuvier.

Dorsal vertebræ 15
Lumbar 6
Dorsal and lumbar together 21
Sacral 5
Total number of vertebræ 26
13 15 14 6 4 5 19 19 19 5 4 4 24 23 23

French
Domestic Boar, from Cuvier.

14
5
19
4
23

Some semi-monstrous breeds deserve notice. From the time of Aristotle to the present time solid-hoofed swine have occasionally been observed in various parts of the world. Although this peculiarity is strongly inherited, it is hardly probable that all the animals with solid hoofs have descended from the same parents; it is more probable that the same peculiarity has reappeared at various times and places. Dr. Struthers has lately described and figured the structure of the feet; in both front and hind feet the distal phalanges of the two greater toes are represented by a single, great, hoof-bearing phalanx; and in the front feet, the middle phalanges are represented by a bone which is single towards the lower end, but bears two separate articulations towards the upper end. From other accounts it appears that an intermediate toe is likewise sometimes superadded.

Another curious anomaly is offered by the appendages, described by M. EudesDeslongchamps as often characterizing the Normandy pigs. These appendages are always attached to the same spot, to the corners of the jaw; they are cylindrical, about three inches in length, covered with bristles, and with a pencil of bristles rising out of a sinus on one side: they have a cartilaginous centre, with two small longitudinal muscles they occur either symmetrically on both sides of the face or on one side alone. Richardson figures them on the gaunt old "Irish Greyhound pig;" and Nathusius states that they occasionally appear in all the long eared races, but are not strictly inherited, for they occur or fail in animals of the same litter.As no wild pigs are known to have analogous appendages, we have at present no reason to suppose that their appearance is due to reversion; and if this be so, we are forced to admit that a somewhat complex, though apparently useless, structure may be suddenly developed without the aid of selection.

It is a remarkable fact that the boars of all domesticated breeds have much shorter tusks than wild boars. Many facts show that with many animals the state of the hair is much affected by exposure to, or protection from, climate; and as we see that the state of the hair and teeth are correlated in Turkish dogs (other analogous facts will be hereafter given), may we not venture to surmise that the reduction of the tusks in the domestic boar is related to his coat of bristles being diminished from living under shelter? On the other hand, as we shall immediately see, the tusks and bristles reappear with feral boars, which are no longer protected from the weather. It is not surprising that the tusks should be more affected than the other teeth; as parts developed to serve as secondary sexual characters are always liable to much variation.

It is a well-known fact that the young of wild European and Indian pigs, for the first six months, are longitudinally banded with light-coloured stripes. This character generally disappears under domestication. The Turkish domestic pigs, however, have striped young, as have those of Westphalia, "whatever may be their hue;" whether these latter pigs belong to the same curly-haired race as the Turkish swine, I do not know. The pigs which have run wild in Jamaica and the semi-feral pigs of New Granada, both those which are black and those which are black with a white band across the stomach, often extending over the back, have resumed this aboriginal character and produce longitudinally-striped young. This is likewise the case, at least occasionally, with the neglected pigs in the Zambesi settlement on the coast of Africa.

The common belief that all domesticated animals, when they run wild, revert completely to the character of their parent-stock, is chiefly founded, as far as I can discover, on feral pigs. But even in this case the belief is not grounded on sufficient evidence; for the two main types, namely, S. scrofa and indicus, have not been distinguished. The young, as we have just seen, reacquire their longitudinal stripes, and the boars invariably reassume their tusks. They revert also in the general shape of their bodies, and in the length of their legs and muzzles, to the state of the wild animal, as might have been expected from the amount of exercise which they are compelled to take in search of food. In Jamaica the feral pigs do not acquire the full size of the European wild boar, "never attaining a greater height than 20 inches at the shoulder." In various countries they reassume their original bristly covering, but in different degrees, dependent on the climate; thus, according to Roulin, the semi-feral pigs in the hot valleys of New Granada are very scantily clothed; whereas, on the Paramos, at the height of 7000 to 8000 feet, they acquire a thick covering of wool lying under the bristles, like that on the truly wild pigs of France. These pigs on the Paramos are small and stunted. The wild boar of India is said to have the bristles at the end of its tail arranged like the plumes of an arrow, whilst the European boar has a simple tuft; and it is a curious fact that many, but not all, of the feral pigs in Jamaica, derived from a Spanish stock, have a plumed tail. With respect to colour, feral pigs generally revert to that of the wild boar; but in certain parts of S. America, as we have seen, some of the semi-feral pigs have a curious white band across their stomachs; and in certain other hot places the pigs are red, and this colour has likewise occasionally been observed in the feral pigs of Jamaica. From these several facts we see that with pigs when feral there is a strong tendency to revert to the wild type; but that this tendency is largely governed by the nature of the climate, amount of exercise, and other causes of change to which they have been subjected.

The last point worth notice is that we have unusually good evidence of breeds of pigs now keeping perfectly true, which have been formed by the crossing of several distinct breeds. The Improved Essex pigs, for instance, breed very true; but there is no doubt that they largely owe their present excellent qualities to crosses originally made by Lord Western with the Neapolitan race, and to subsequent crosses with the Berkshire breed (this also having been improved by Neapolitan crosses), and likewise, probably, with the Sussex breed. In breeds thus formed by complex crosses, the most careful and unremitting selection during many generations has been found to be indispensable. Chiefly in consequence of so much crossing, some well-known breeds have undergone rapid changes; thus, according to Nathusius, the Berkshire breed of 1780 is quite different from that of 1810; and, since this latter period, at least two distinct forms have borne the same name.

CATTLE.

Domestic cattle are certainly the descendants of more than one wild form, in the same manner as has been shown to be the case with our dogs and pigs. Naturalists have generally made two main divisions of cattle: the humped kinds inhabiting tropical countries, called in India Zebus, to which the specific name of Bos indicus has been given; and the common non-humped cattle, generally included under the name of Bos taurus. The humped cattle were domesticated, as may be seen on the Egyptian monuments, at least as early as the twelfth dynasty, that is 2100 B.C. They differ from common cattle in various osteological characters, even in a greater degree, according to Rütimeyer, than do the fossil and prehistoric European species, namely, Bos primigenius and longifrons, from each other. They differ, also, as Mr. Blyth, who has particularly attended to this subject, remarks, in general configuration, in the shape of their ears, in the point where the dewlap commences, in the typical curvature of their horns, in their manner of carrying their heads when at rest, in their ordinary variations of colour, especially in the frequent presence of "nilgau-like markings on their feet," and "in the one being born with teeth protruding through the jaws, and the other not so." They have different habits, and their voice is entirely different. The humped cattle in India "seldom seek shade, and never go into the water and there stand knee-deep, like the cattle of Europe." They have run wild in parts of Oude and Rohilcund, and can maintain themselves in a region infested by tigers. They have given rise to many races differing greatly in size, in the presence of one or two humps, in length of horns, and other respects. Mr. Blyth sums up emphatically that the humped and humpless cattle must be considered as distinct species. When we consider the number of points in external structure and habits, independently of important osteological differences, in which they differ from each other; and that many of these points are not likely to have been affected by domestication, there can hardly be a doubt, notwithstanding the adverse opinion of some naturalists, that the humped and non-humped cattle must be ranked as specifically distinct.

The European breeds of humpless cattle are numerous. Professor Low enumerates 19 British breeds, only a few of which are identical with those on the Continent. Even the small Channel islands of Guernsey, Jersey, and Alderney possess their own sub-breeds; and these again differ from the cattle of the other British islands, such as Anglesea, and the western isles of Scotland. Desmarest, who paid attention to the subject, describes 15 French races, excluding sub-varieties and those imported from other countries. In other parts of Europe there are several distinct races, such as the pale-coloured Hungarian cattle, with their light and free step, and enormous horns sometimes measuring above five feet from tip to tip: the Podolian cattle also are remarkable from the height of their forequarters. In the most recent work on Cattle, engravings are given of fifty-five European breeds; it is, however, probable that several of these differ very little from each other, or are merely synonyms. It must not be supposed that numerous breeds of cattle exist only in long-civilised countries, for we shall presently see that several kinds are kept by the savages of Southern Africa.

With respect to the parentage of the several European breeds, we already know much from Nilsson's Memoir, and more especially from Rütimeyer's works and those of Boyd Dawkins. Two or three species or forms of Bos, closely allied to still living domestic races, have been found in the more recent tertiary deposits or amongst prehistoric remains in Europe. Following Rütimeyer, we have:—

Bos primigenius. This magnificent, well known species was domesticated in Switzerland during the Neolithic period; even at this early period it varied a little, having apparently been crossed with other races. Some of the larger races on the Continent, as the Friesland, etc., and the Pembroke race in England, closely resemble in essential structure B. primigenius, and no doubt are its descendants. This is likewise the opinion of Nilsson. Bos primigenius existed as a wild animal in Cæsar's time, and is now semi-wild, though much degenerated in size, in the park of Chillingham; for I am informed by Professor Rütimeyer, to whom Lord Tankerville sent a skull, that the Chillingham cattle are less altered from the true primigenius type than any other known breed.

Bos trochoceros. This form is not included in the three species above mentioned, for it is now considered by Rütimeyer to be the female of an early domesticated form of B. primigenius, and as the progenitor of his frontosus race. I may add that specific names have been given to four other fossil oxen, now believed to be identical with B. primigenius.

Bos longifrons (or brachyceros) of Owen.—This very distinct species was of small size, and had a short body with fine legs. According to Boyd