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 ones 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 Mr. 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 vertebræ 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 vertebræ, 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 a 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 also 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 even breed with their own females, so that the negative results obtained from crosses are of no value, and the antipathy of wild animals of different 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 ensuring 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 276 of his work:—
“First crosses between forms sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the two most careful experimentalists who have ever lived have come to diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible of favourable and unfavourable conditions. The degree of sterility does not strictly follow systematic affinity, but is governed by several curious and complex laws. It is generally different, and sometimes widely different, in reciprocal crosses between the same two species. It is not always equal in degree in a first cross, and in the hybrid produced from this cross.
“In the same manner as in grafting trees, the capacity of one species or variety to take on another is incidental on generally unknown differences in their vegetative systems, so in crossing, the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. There is no more reason to think that species have been specially endowed with various degrees of sterility to prevent them crossing and breeding in nature, than to think that trees have been specially endowed with various and somewhat analogous degrees of difficulty in being grafted together, in order to prevent them becoming inarched in our forests.
“The sterility of first crosses between pure species, which have their reproductive systems perfect, seems to depend on several circumstances; in some cases largely on the early death of the embryo. The sterility of hybrids which have their reproductive systems imperfect, and which have had this system and their whole organization disturbed by being compounded of two distinct species, seems closely allied to that sterility which so frequently affects pure species when their natural conditions of life have been disturbed. This view is supported by a parallelism of another kind; namely, that the crossing of forms only slightly different is favourable to the vigour and fertility of the offspring; and that slight changes in the conditions of life are apparently favourable to the vigour and fertility of all organic beings. It is not surprising that the degree of difficulty in uniting two species, and the degree of sterility of their hybrid offspring should generally correspond, though due to distinct causes; for both depend on the amount of difference of some kind between the species which are crossed. Nor is it surprising that the facility of effecting a first cross, the fertility of hybrids produced from it, and the capacity of being grafted together—though this latter capacity evidently depends on widely different circumstances—should all run to a certain extent parallel with the systematic affinity of the forms which are subjected to experiment; for systematic affinity attempts to express all kinds of resemblance between all species.
“First crosses between forms known to be varieties, or sufficiently alike to be considered as varieties, and their mongrel offspring, are very generally, but not quite universally, fertile. Nor is this nearly general and perfect fertility surprising, when we remember how liable we are to argue in a circle with respect to varieties in a state of nature; and when we remember that the greater number of varieties have been produced under domestication by the selection of mere external differences, and not of differences in the reproductive system. In all other respects, excluding fertility, there is a close general resemblance between hybrids and mongrels” (pp. 276-8).