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    The Origin of Species

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    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.

     


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