On the Origin of Species by Means of Natural Selection
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The doctrine of the origin of our several domestic races from several aboriginal stocks, has been carried to an absurd extreme by some authors. They believe that every race which breeds true, let the distinctive characters be ever so slight, has had its wild prototype. At this rate there must have existed at least a score of species of wild cattle, as many sheep, and several goats in Europe alone, and several even within Great Britain. One author believes that there formerly existed in Great Britain eleven wild species of sheep peculiar to it! When we bear in mind that Britain has now hardly one peculiar mammal, and France but few distinct from those of Germany and conversely, and so with Hungary, Spain, &c., but that each of these kingdoms possesses several peculiar breeds of cattle, sheep, &c., we must admit that many domestic breeds have originated in Europe; for whence could they have been derived, as these several countries do not possess a number of peculiar species as distinct parent-stocks? So it is in India. Even in the case of the domestic dogs of the whole world, which I fully admit have probably descended from several wild species, I cannot doubt that there has been an immense amount of inherited variation. Who can believe that animals closely resembling the Italian greyhound, the bloodhound, the bull-dog, or Blenheim spaniel, &c.--so unlike all wild Canidae--ever existed freely in a state of nature? It has often been loosely said that all our races of dogs have been produced by the crossing of a few aboriginal species; but by crossing we can get only forms in some degree intermediate between their parents; and if we account for our several domestic races by this process, we must admit the former existence of the most extreme forms, as the Italian greyhound, bloodhound, bull-dog, &c., in the wild state. Moreover, the possibility of making distinct races by crossing has been greatly exaggerated. There can be no doubt that a race may be modified by occasional crosses, if aided by the careful selection of those individual mongrels, which present any desired character; but that a race could be obtained nearly intermediate between two extremely different races or species, I can hardly believe. Sir J. Sebright expressly experimentised for this object, and failed. The offspring from the first cross between two pure breeds is tolerably and sometimes (as I have found with pigeons) extremely uniform, and everything seems simple enough; but when these mongrels are crossed one with another for several generations, hardly two of them will be alike, and then the extreme difficulty, or rather utter hopelessness, of the task becomes apparent. Certainly, a breed intermediate between two very distinct breeds could not be got without extreme care and long-continued selection; nor can I find a single case on record of a permanent race having been thus formed.

On the Breeds of the Domestic Pigeon. -- 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 considerably 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 and strictly 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 very long beak, has a very short and very 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 very 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 oesophagus. 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, much 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 members of the great pigeon family; and 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 have been specified.

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 number of the caudal and sacral vertebrae vary; 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 oesophagus; the development and abortion of the oil-gland; the number of the primary wing and caudal feathers; the relative length of wing and tail to each other and to the body; the relative length of leg and of the feet; the number of scutellae 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 differs remarkably; as does in some breeds the voice and disposition. Lastly, in certain breeds, the males and females have come to differ to a slight degree from each other.

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, I think, be ranked by him as well-defined species. Moreover, I do not believe that any ornithologist would 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 might have called them, could be shown him.

Great as the differences are between the breeds of pigeons, I am fully convinced that the common opinion of naturalists is correct, namely, that all have 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, not breeding or willingly perching 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 very 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 to me 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 ever become wild or feral, though the dovecot-pigeon, which is the rock-pigeon in a very slightly altered state, has become feral in several places. Again, all recent experience shows that it is most difficult to get any wild animal 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-civilized man, as to be quite prolific under confinement.

An argument, as it seems to me, of great weight, and applicable in several other cases, is, that the above-specified breeds, though agreeing generally in constitution, habits, voice, colouring, and in most parts of their structure, with the wild rock-pigeon, yet are certainly highly abnormal in other parts of their structure: we may look in vain throughout the whole great family of Columbidae 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-civilized 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 seem to me improbable in the highest degree.

Some facts in regard to the colouring of pigeons well deserve consideration. The rock-pigeon is of a slaty-blue, and has a white rump (the Indian sub-species, C. intermedia of Strickland, having it bluish); the tail has a terminal dark bar, with the bases of the outer feathers externally edged with white; the wings have two black bars; some semi-domestic breeds and some apparently 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 two birds belonging to two distinct breeds are crossed, neither of which is blue or has any of the above-specified marks, the mongrel offspring are very apt suddenly to acquire these characters; for instance, I crossed some uniformly white fantails with some uniformly black barbs, and they produced mottled brown and black birds; these I again crossed together, and one grandchild of the pure white fantail and pure black barb was of as beautiful a blue colour, with the white rump, 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 have descended from the rock-pigeon. But if we deny this, we must make one of the two following highly improbable suppositions. Either, firstly, 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 a dozen or twenty generations, for we know of no fact countenancing the belief that the child ever reverts to some one ancestor, removed by a greater number of generations. In a breed which has been crossed only once with some distinct breed, the tendency to reversion to any character derived from such 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 with a distinct breed, and there is a tendency in both parents to revert to a character, which has been 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 are often confounded in treatises on inheritance.

Lastly, the hybrids or mongrels from between all the domestic breeds of pigeons are perfectly fertile. I can state this from my own observations, purposely made on the most distinct breeds. Now, it is difficult, perhaps impossible, to bring forward one case of the hybrid offspring of two animals clearly distinct being themselves perfectly fertile. Some authors believe that long-continued domestication eliminates this strong tendency to sterility: from the history of the dog I think there is some probability in this hypothesis, if applied to species closely related together, though it is unsupported by a single experiment. But to extend the hypothesis 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, seems to me rash in the extreme.

From these several reasons, namely, the improbability of man having formerly got seven or eight supposed species of pigeons to breed freely under domestication; these supposed species being quite unknown in a wild state, and their becoming nowhere feral; these species having very abnormal characters in certain respects, as compared with all other Columbidae, though so like in most other respects to the rock-pigeon; the blue colour and various marks occasionally appearing in all the breeds, both when kept pure and when crossed; the mongrel offspring being perfectly fertile;--from these several reasons, taken together, I can feel no doubt that all our domestic breeds have descended from the Columba livia with its geographical sub-species.

In favour of this view, I may add, firstly, that C. livia, or the rock-pigeon, 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, although an English carrier or short-faced tumbler differs immensely in certain characters from the rock-pigeon, yet by comparing the several sub-breeds of these breeds, more especially those brought from distant countries, we can make an almost perfect series between the extremes of structure. Thirdly, those characters which are mainly distinctive of each breed, 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, are in each breed eminently variable; and the explanation of this fact will be obvious when we come to 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 Aegyptian 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 20,000 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 be obvious when we treat of Selection. We shall then, also, see how it is that the 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.

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, knowing well how true they bred, I felt fully as much difficulty in believing that they could ever have descended from a common parent, as any naturalist could in coming to a similar conclusion in regard to the many species of finches, or other large groups of birds, in nature. One circumstance has struck me much; namely, that all the breeders of the various domestic animals and the cultivators of plants, with whom I have ever 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, 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 have 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?

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