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Charles Robert Darwin (1809–1882).  Origin of Species.
The Harvard Classics.  1909–14.
 
I. Variation under Domestication
 
Breeds of the Domestic Pigeon, Their Differences and Origin
 
 
BELIEVING that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world, more especially by the Hon. W. Elliot from India, and by the Hon. C. Murray from Persia. Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerable antiquity. I have associated with several eminent fanciers, and have been permitted to join two of the London Pigeon Clubs. The diversity of the breeds is something astonishing. Compare the English carrier and the short-faced tumbler, and see the wonderful difference in their beaks, entailing corresponding differences in their skulls. The carrier, more especially the male bird, is also remarkable from the wonderful development of the carunculated skin about the head; and this is accompanied by greatly elongated eyelids, very large external orifices to the nostrils, and a wide gape of mouth. The short-faced tumbler has a beak in outline almost like that of a finch; and the common tumbler has the singular inherited habit of flying at a great height in a compact flock, and tumbling in the air head over heels. The runt is a bird of great size, with long massive beak and large feet; some of the sub-breeds of runts have very long necks, others very long wings and tails, others singularly short tails. The barb is allied to the carrier, but, instead of a long beak has a very short and broad one. The pouter has a much elongated body, wings, and legs; and its enormously developed crop, which it glories in inflating, may well excite astonishment and even laughter. The turbit has a short and conical beak, with a line of reversed feathers down the breast; and it has the habit of continually expanding slightly, the upper part of the œsophagus. The Jacobin has the feathers so much reversed along the back of the neck that they form a hood; and it has, proportionally to its size, elongated wing and tail feathers. The trumpeter and laugher, as their names express, utter a very different coo from the other breeds. The fantail has thirty or even forty tail-feathers, instead of twelve or fourteen—the normal number in all the members of the great pigeon family: these feathers are kept expanded, and are carried so erect, that in good birds the head and tail touch: the oil-gland is quite aborted. Several other less distinct breeds might be specified.  1
  In the skeletons of the several breeds, the development of the bones of the face in length and breadth and curvature differs enormously. The shape, as well as the breadth and length of the ramus of the lower jaw, varies in a highly remarkable manner. The caudal and sacral vertebræ vary in number; as does the number of the ribs, together with their relative breadth and the presence of processes. The size and shape of the apertures in the sternum are highly variable; so is the degree of divergence and relative size of the two arms of the furcula. The proportional width of the gape of mouth, the proportional length of the eyelids, of the orifice of the nostrils, of the tongue (not always in strict correlation with the length of beak), the size of the crop and of the upper part of the œsophagus; the development and abortion of the oil-gland; the number of the primary wing and caudal feathers; the relative length of the wing and tail to each other and to the body; the relative length of the leg and foot; the number of scutellæ on the toes, the development of skin between the toes, are all points of structure which are variable. The period at which the perfect plumage is acquired varies, as does the state of the down with which the nestling birds are clothed when hatched. The shape and size of the eggs vary. The manner of flight, and in some breeds the voice and disposition, differ remarkably. Lastly, in certain breeds, the males and females have come to differ in a slight degree from each other.  2
  Altogether at least a score of pigeons might be chosen, which, if shown to an ornithologist, and he were told that they were wild birds, would certainly be ranked by him as well-defined species. Moreover, I do not believe that any ornithologist would in this case place the English carrier, the short-faced tumbler, the runt, the barb, pouter, and fantail in the same genus; more especially as in each of these breeds several truly inherited sub-breeds, or species, as he would call them, could be shown him.  3
  Great as are the differences between the breeds of the pigeon, I am fully convinced that the common opinion of naturalists is correct, namely, that all are descended from the rock pigeon (Columba livia), including under this term several geographical races or sub-species, which differ from each other in the most trifling respects. As several of the reasons which have led me to this belief are in some degree applicable in other cases, I will here briefly give them. If the several breeds are not varieties, and have not proceeded from the rock pigeon, they must have descended from at least seven or eight aboriginal stocks; for it is impossible to make the present domestic breeds by the crossing of any lesser number: how, for instance, could a pouter be produced by crossing two breeds unless one of the parent-stocks possessed the characteristic enormous crop? The supposed aboriginal stocks must all have been rock pigeons, that is, they did not breed or willingly perch on trees. But besides C. livia, with its geographical sub-species, only two or three other species of rock pigeons are known; and these have not any of the characters of the domestic breeds. Hence the supposed aboriginal stocks must either still exist in the countries where they were originally domesticated, and yet be unknown to ornithologists; and this, considering their size, habits, and remarkable characters, seems improbable; or they must have become extinct in the wild state. But birds breeding on precipices, and good fliers, are unlikely to be exterminated; and the common rock pigeon, which has the same habits with the domestic breeds, has not been exterminated even on several of the smaller British islets, or on the shores of the Mediterranean. Hence the supposed extermination of so many species having similar habits with the rock pigeon seems a very rash assumption. Moreover, the several above-named domesticated breeds have been transported to all parts of the world, and, therefore, some of them must have been carried back again into their native country; but not one has become wild or feral, though the dovecot pigeon, which is the rock pigeon in very slightly altered state, has become feral in several places. Again, all recent experience shows that it is difficult to get wild animals to breed freely under domestication, yet on the hypothesis of the multiple origin of our pigeons, it must be assumed that at least seven or eight species were so thoroughly domesticated in ancient times by half-civilised man, as to be quite prolific under confinement.  4
  An argument of great weight, and applicable in several other cases, is, that the above-specified breeds, though agreeing generally with the wild rock pigeon in constitution, habits, voice, colouring, and in most parts of their structure, yet are certainly highly abnormal in other parts; we may look in vain through the whole great family of Columbidæ for a beak like that of the English carrier, or that of the short-faced tumbler, or barb; for reversed feathers like those of the Jacobin; for a crop like that of the pouter; for tail-feathers like those of the fantail. Hence it must be assumed not only that half-civilised man succeeded in thoroughly domesticating several species, but that he intentionally or by chance picked out extraordinarily abnormal species; and further, that these very species have since all become extinct or unknown. So many strange contingencies are improbable in the highest degree.  5
  Some facts in regard to the colouring of pigeons well deserve consideration. The rock pigeon is of a slaty-blue, with white loins; but the Indian sub-species, C. intermedia of Strickland, has this part bluish. The tail has a terminal dark bar, with the outer feathers externally edged at the base with white. The wings have two black bars. Some semi-domestic breeds, and some truly wild breeds, have, besides the two black bars, the wings chequered with black. These several marks do not occur together in any other species of the whole family. Now, in every one of the domestic breeds, taking thoroughly well-bred birds, all the above marks, even to the white edging of the outer tail-feathers, sometimes concur perfectly developed. Moreover, when birds belonging to two or more distinct breeds are crossed, none of which are blue or have any of the above-specified marks, the mongrel offspring are very apt suddenly to acquire these characters. To give one instance out of several which I have observed:—I crossed some white fantails, which breed very true, with some black barbs—and it so happens that blue varieties of barbs are so rare that I never heard of an instance in England; and the mongrels were black, brown, and mottled. I also crossed a barb with a spot, which is a white bird with a red tail and red spot on the forehead, and which notoriously breeds very true; the mongrels were dusky and mottled. I then crossed one of the mongrel barb-fantails with a mongrel barb-spot, and they produced a bird of as beautiful a blue colour, with the white loins, double black wing-bar, and barred and white-edged tail-feathers, as any wild rock pigeon! We can understand these facts, on the well-known principle of reversion to ancestral characters, if all the domestic breeds are descended from the rock pigeon. But if we deny this, we must make one of the two following highly improbable suppositions. Either, first, that all the several imagined aboriginal stocks were coloured and marked like the rock pigeon, although no other existing species is thus coloured and marked, so that in each separate breed there might be a tendency to revert to the very same colours and markings. Or, secondly, that each breed, even the purest, has within a dozen, or at most within a score, of generations, been crossed by the rock pigeon: I say within dozen or twenty generations, for no instance is known of crossed descendants reverting to an ancestor of foreign blood, removed by a greater number of generations. In a breed which has been crossed only once, the tendency to revert to any character derived from such a cross will naturally become less and less, as in each succeeding generation there will be less of the foreign blood; but when there has been no cross, and there is a tendency in the breed to revert to a character which was lost during some former generation, this tendency, for all that we can see to the contrary, may be transmitted undiminished for an indefinite number of generations. These two distinct cases of reversion are often confounded together by those who have written on inheritance.  6
  Lastly, the hybrids or mongrels from between all the breeds of the pigeon are perfectly fertile, as I can state from my own observations, purposely made, on the most distinct breeds. Now, hardly any cases have been ascertained with certainty of hybrids from two quite distinct species of animals being perfectly fertile. Some authors believe that long-continued domestication eliminates this strong tendency to sterility in species. From the history of the dog, and of some other domestic animals, this conclusion is probably quite correct, if applied to species closely related to each other. But to extend it so far as to suppose that species, aboriginally as distinct as carriers, tumblers, pouters, and fantails now are, should yield offspring perfectly fertile inter se, would be rash in the extreme.  7
  From these several reasons, namely,—the improbability of man having formerly made seven or eight supposed species of pigeons to breed freely under domestication;—these supposed species being quite unknown in a wild state, and their not having become anywhere feral;—these species presenting certain very abnormal characters, as compared with all other Columbidæ, though so like the rock pigeon in most respects;—the occasional re-appearance of the blue colour and various black marks in all the breeds, both when kept pure and when crossed;—and lastly, the mongrel offspring being perfectly fertile;—from these several reasons taken together, we may safely conclude that all our domestic breeds are descended from the rock pigeon or Columba livia with its geographical sub-species.  8
  In favour of this view, I may add, firstly, that the wild C. livia has been found capable of domestication in Europe and in India; and that it agrees in habits and in a great number of points of structure with all the domestic breeds. Secondly, that, although an English carrier or a short-faced tumbler differs immensely in certain characters from the rock pigeon, yet that, by comparing the several sub-breeds of these two races, more especially those brought from distant countries, we can make, between them and the rock pigeon, an almost perfect series; so we can in some other cases, but not with all the breeds. Thirdly, those characters which are mainly distinctive of each breed are in each eminently variable, for instance the wattle and length of beak of the carrier, the shortness of that of the tumbler, and the number of tail-feathers in the fantail; and the explanation of this fact will be obvious when we treat of Selection. Fourthly, pigeons have been watched and tended with the utmost care, and loved by many people. They have been domesticated for thousands of years in several quarters of the world; the earliest known record of pigeons is in the fifth Ægyptian dynasty, about 3000 B.C., as was pointed out to me by Professor Lepsius; but Mr. Birch informs me that pigeons are given in a bill of fare in the previous dynasty. In the time of the Romans, as we hear from Pliny, immense prices were given for pigeons; “nay, they are come to this pass, that they can reckon up their pedigree and race.” Pigeons were much valued by Akber Khan in India, about the year 1600; never less than 90,000 pigeons were taken with the court. “The monarchs of Iran and Turan sent him some very rare birds”; and continues the courtly historian, “His Majesty by crossing the breeds, which method was never practised before, has improved them astonishingly.” About this same period the Dutch were as eager about pigeons as were the old Romans. The paramount importance of these considerations in explaining the immense amount of variation which pigeons have undergone, will likewise be obvious when we treat of Selection. We shall then, also, see how it is that the several breeds so often have a somewhat monstrous character. It is also a most favourable circumstance for the production of distinct breeds, that male and female pigeons can be easily mated for life; and thus different breeds can be kept together in the same aviary.  9
  I have discussed the probable origin of domestic pigeons at some, yet quite insufficient, length; because when I first kept pigeons and watched the several kinds, well knowing how truly they breed, I felt fully as much difficulty in believing that since they had been domesticated they had all proceeded from a common parent, as any naturalist could in coming to a similar conclusion in regard to the many species of finches, or other groups of birds, in nature. One circumstance has struck me much; namely, that nearly all the breeders of the various domestic animals and the cultivators of plants, with whom I have conversed, or whose treatises I have read, are firmly convinced that the several breeds to which each has attended, are descended from so many aboriginally distinct species. Ask, as I have asked, a celebrated raiser of Hereford cattle, whether his cattle might not have descended from long-horns, or both from a common parent-stock, and he will laugh you to scorn. I have never met a pigeon, or poultry, or duck, or rabbit fancier, who was not fully convinced that each main breed was descended from a distinct species. Van Mons, in his treatise on pears and apples, shows how utterly he disbelieves that the several sorts, for instance a Ribston-pippin or Codlin-apple, could ever have proceeded from the seeds of the same tree. Innumerable other examples could be given. The explanation, I think, is simple: from long-continued study they are strongly impressed with the differences between the several races; and though they well know that each race varies slightly, for they win their prizes by selecting such slight differences, yet they ignore all general arguments, and refuse to sum up in their minds slight differences accumulated during many successive generations. May not those naturalists who, knowing far less of the laws of inheritance than does the breeder, and knowing no more than he does of the intermediate links in the long lines of descent, yet admit that many of our domestic races are descended from the same parents—may they not learn a lesson of caution, when they deride the idea of species in a state of nature being lineal descendants of other species?  10
 

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