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- THE ORIGIN OF SPECIES - 3/7 -


offspring was always pure Ancon. Colonel Humphreys, in fact, states that he was acquainted with only "one questionable case of a contrary nature." Here, then, is a remarkable and well-established instance, not only of a very distinct race being established 'per saltum', but of that race breeding "true" at once, and showing no mixed forms, even when crossed with another breed.

[footnote] *Colonel Humphreys' statements are exceedingly explicit on this point:--"When an Ancon ewe is impregnated by a common ram, the increase resembles wholly either the ewe or the ram. The increase of the common ewe impregnated by an Ancon ram follows entirely the one or the other, without blending any of the distinguishing and essential peculiarities of both. Frequent instances have happened where common ewes have had twins by Ancon rams, when one exhibited the complete marks and features of the ewe, the other of the ram. The contrast has been rendered singularly striking, when one short-legged and one long-legged lamb, produced at a birth, have been seen sucking the dam at the same time."--'Philosophical Transactions', 1813, Pt. I. pp. 89, 90.

By taking care to select Ancons of both sexes, for breeding from, it thus became easy to establish an extremely well-marked race; so peculiar that, even when herded with other sheep, it was noted that the Ancons kept together. And there is every reason to believe that the existence of this breed might have been indefinitely protracted; but the introduction of the Merino sheep, which were not only very superior to the Ancons in wool and meat, but quite as quiet and orderly, led to the complete neglect of the new breed, so that, in 1813, Colonel Humphreys found it difficult to obtain the specimen, whose skeleton was presented to Sir Joseph Banks. We believe that, for many years, no remnant of it has existed in the United States.

Gratio Kelleia was not the progenitor of a race of six-fingered men, as Seth Wright's ram became a nation of Ancon sheep, though the tendency of the variety to perpetuate itself appears to have been fully as strong in the one case as in the other. And the reason of the difference is not far to seek. Seth Wright took care not to weaken the Ancon blood by matching his Ancon ewes with any but males of the same variety, while Gratio Kelleia's sons were too far removed from the patriarchal times to intermarry with their sisters; and his grandchildren seem not to have been attracted by their six-fingered cousins. In other words, in the one example a race was produced, because, for several generations, care was taken to 'select' both parents of the breeding stock from animals exhibiting a tendency to vary in the same condition; while, in the other, no race was evolved, because no such selection was exercised. A race is a propagated variety; and as, by the laws of reproduction, offspring tend to assume the parental forms, they will be more likely to propagate a variation exhibited by both parents than that possessed by only one.

There is no organ of the body of an animal which may not, and does not, occasionally, vary more or less from the normal type; and there is no variation which may not be transmitted and which, if selectively transmitted, may not become the foundation of a race. This great truth, sometimes forgotten by philosophers, has long been familiar to practical agriculturists and breeders; and upon it rest all the methods of improving the breeds of domestic animals, which, for the last century, have been followed with so much success in England. Colour, form, size, texture of hair or wool, proportions of various parts, strength or weakness of constitution, tendency to fatten or to remain lean, to give much or little milk, speed, strength, temper, intelligence, special instincts; there is not one of these characters whose transmission is not an every-day occurrence within the experience of cattle-breeders, stock-farmers, horse-dealers, and dog and poultry fanciers. Nay, it is only the other day that an eminent physiologist, Dr. Brown-Sequard, communicated to the Royal Society his discovery that epilepsy, artificially produced in guinea-pigs, by a means which he has discovered, is transmitted to their offspring.

But a race, once produced, is no more a fixed and immutable entity than the stock whence it sprang; variations arise among its members, and as these variations are transmitted like any others, new races may be developed out of the pre-existing one 'ad infinitum', or, at least, within any limit at present determined. Given sufficient time and sufficiently careful selection, and the multitude of races which may arise from a common stock is as astonishing as are the extreme structural differences which they may present. A remarkable example of this is to be found in the rock-pigeon, which Dr. Darwin has, in our opinion, satisfactorily demonstrated to be the progenitor of all our domestic pigeons, of which there are certainly more than a hundred well-marked races. The most noteworthy of these races are, the four great stocks known to the "fancy" as tumblers, pouters, carriers, and fantails; birds which not only differ most singularly in size, colour, and habits, but in the form of the beak and of the skull: in the proportions of the beak to the skull; in the number of tail-feathers; in the absolute and relative size of the feet; in the presence or absence of the uropygial gland; in the number of vertebrae in the back; in short, in precisely those characters in which the genera and species of birds differ from one another.

And it is most remarkable and instructive to observe, that none of these races can be shown to have been originated by the action of changes in what are commonly called external circumstances, upon the wild rock-pigeon. On the contrary, from time immemorial, pigeon-fanciers have had essentially similar methods of treating their pets, which have been housed, fed, protected and cared for in much the same way in all pigeonries. In fact, there is no case better adapted than that of the pigeons to refute the doctrine which one sees put forth on high authority, that "no other characters than those founded on the development of bone for the attachment of muscles" are capable of variation. In precise contradiction of this hasty assertion, Mr. Darwin's researches prove that the skeleton of the wings in domestic pigeons has hardly varied at all from that of the wild type; while, on the other hand, it is in exactly those respects, such as the relative length of the beak and skull, the number of the vertebrae, and the number of the tail-feathers, in which muscular exertion can have no important influence, that the utmost amount of variation has taken place.

We have said that the following out of the properties exhibited by physiological species would lead us into difficulties, and at this point they begin to be obvious; for if, as the result of spontaneous variation and of selective breeding, the progeny of a common stock may become separated into groups distinguished from one another by constant, not sexual, morphological characters, it is clear that the physiological definition of species is likely to clash with the morphological definition. No one would hesitate to describe the pouter and the tumbler as distinct species, if they were found fossil, or if their skins and skeletons were imported, as those of exotic wild birds commonly are--and without doubt, if considered alone, they are good and distinct morphological species. On the other hand, they are not physiological species, for they are descended from a common stock, the rock-pigeon.

Under these circumstances, as it is admitted on all sides that races occur in Nature, how are we to know whether any apparently distinct animals are really of different physiological species, or not, seeing that the amount of morphological difference is no safe guide? Is there any test of a physiological species? The usual answer of physiologists is in the affirmative. It is said that such a test is to be found in the phenomena of hybridization--in the results of crossing races, as compared with the results of crossing species.

So far as the evidence goes at present, individuals, of what are certainly known to be mere races produced by selection, however distinct they may appear to be, not only breed freely together, but the offspring of such crossed races are only perfectly fertile with one another. Thus, the spaniel and the greyhound, the dray-horse and the Arab, the pouter and the tumbler, breed together with perfect freedom, and their mongrels, if matched with other mongrels of the same kind, are equally fertile.

On the other hand, there can be no doubt that the individuals of many natural species are either absolutely infertile if crossed with individuals of other species, or, if they give rise to hybrid offspring, the hybrids so produced are infertile when paired together. The horse and the ass, for instance, if so crossed, give rise to the mule, and there is no certain evidence of offspring ever having been produced by a male and female mule. The unions of the rock-pigeon and the ring-pigeon appear to be equally barren of result. Here, then, says the physiologist, we have a means of distinguishing any two true species from any two varieties. If a male and a female, selected from each group, produce offspring, and that offspring is fertile with others produced in the same way, the groups are races and not species. If, on the other hand, no result ensues, or if the offspring are infertile with others produced in the same way, they are true physiological species. The test would be an admirable one, if, in the first place, it were always practicable to apply it, and if, in the second, it always yielded results susceptible of a definite interpretation. Unfortunately, in the great majority of cases, this touchstone for species is wholly inapplicable.

The constitution of many wild animals is so altered by confinement that they will not breed even with their own females, so that the negative results obtained from crosses are of no value; and the antipathy of wild animals of the same species for one another, or even of wild and tame members of the same species, is ordinarily so great, that it is hopeless to look for such unions in Nature. The hermaphrodism of most plants, the difficulty in the way of insuring the absence of their own, or the proper working of other pollen, are obstacles of no less magnitude in applying the test to them. And, in both animals and plants, is superadded the further difficulty, that experiments must be continued over a long time for the purpose of ascertaining the fertility of the mongrel or hybrid progeny, as well as of the first crosses from which they spring.

Not only do these great practical difficulties lie in the way of applying the hybridization test, but even when this oracle can be questioned, its replies are sometimes as doubtful as those of Delphi. For example, cases are cited by Mr. Darwin, of plants which are more fertile with the pollen of another species than with their own; and there are others, such as certain 'fuci', whose male element will fertilize the ovule of a plant of distinct species, while the males of the latter species are ineffective with the females of the first. So that, in the last-named instance, a physiologist, who should cross the two species in one way, would decide that they were true species; while another, who should cross them in the reverse way, would, with equal justice, according to the rule, pronounce them to be mere races. Several plants, which there is great reason to believe are mere varieties, are almost sterile when crossed; while both animals and plants, which have always been regarded by naturalists as of distinct species, turn out, when the test is applied, to be perfectly fertile. Again, the sterility or fertility of crosses seems to bear no relation to the structural resemblances or differences of the members of any two groups.

Mr. Darwin has discussed this question with singular ability and circumspection, and his conclusions are summed up as follows, at page


THE ORIGIN OF SPECIES - 3/7

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