Cross-infertility in Domesticated Cattle.—Mr. J. W. Crompton, who has had a large experience as a professional cattle-breeder, writes to me (March 2, 1887)—
"That form of barrenness, very common in some districts, which makes heifers become what are called 'bullers'—that is, irregularly in 'season,' wild, and failing to conceive—is certainly produced by excess of iron in their drinking-water, and I suspect also by a deficiency of potash in the soil."
He also informs me that pure white beasts of either sex are so well known by experienced breeders to be comparatively infertile together, that they are never used for breeding purposes, so that "in some parts of the country, where a tendency to sterility had become so confirmed in the white race that they utterly died out," only the coloured breeds are now to be found. He goes on to say that if "a lot of white heifers were put to a lot of white bulls, I think you would probably get a fertile breed of pure white cattle.... I think, in short, that domestication has produced just what your theory suggests, a new variety inclined to prove sterile with its parent stock."
Commenting on the origin of domesticated cattle, Professor Oscar Schmidt remarks (Doctrine of Descent, p. 139)—
"Rütimeyer's minute researches on domestic cattle have shown that, in Europe at least, three well-defined species of the diluvial period have contributed to their formation—Bos primigenius, longifrons, and frontosus. These species once lived geographically separate, but contemporaneously; and they and their specific peculiarities have perished, to rise again in our domestic races. These races breed together with unqualified fertility. In the form of skull and horns they recall one or other of the extinct species; but collectively they constitute a new main species. That from their various breeds, the three or any one of the aboriginal species would ever emerge in a state of pristine purity, would be an utterly ludicrous assertion."
Now, seeing that these "aboriginal species," although living "contemporaneously," were "geographically separate," we can well understand that their divergence of type from a common ancestor did not require, as a condition to their divergence, that any cross-sterility should have arisen between them. The geographical isolation was enough to secure immunity from mutual intercrossing, and therefore, as our present theory would have expected as probable, morphological divergence occurred without any corresponding physiological divergence, as must almost certainly have been the case if such polytypic evolution had occurred on a common area. Indeed, one of the two lines of experimental verification of our theory consists in selecting cases where nearly allied species are separated by geographical barriers, and proving that, in such cases, there is no cross-sterility.
Fertility of Domesticated Varieties.—Some writers have sought to explain the contrast between domesticated varieties and natural species in respect of fertility when crossed, by the consideration that it is only those natural species which have proved themselves so far flexible as to continue fertile under changed conditions of life that can have ever allowed themselves to become domesticated. But although this condition may well serve to explain the unimpaired fertility under domestication of such species as for this very reason have ever become domesticated, I fail to see how it explains the further and altogether different fact, that this fertility continues unimpaired between all the newly differentiated morphological types which have been derived from the original specific type. It is one thing that this type should continue fertile after domestication: it is quite another thing that fertility should continue as between all its modified descendants, even although the amount of modification may extend much further than that which usually obtains between different natural species.
Testing for Cross-infertility among varieties growing on the same area is a much more crucial line of verification than testing for unimpaired fertility between allied species which occupy different areas, because while in the former case we are dealing with "incipient species" with a view to ascertaining whether the divergence which they have already undergone is accompanied by physiological isolation, in the latter case we can never be sure that two allied species, which are now widely disconnected geographically, have always been so disconnected. They may both have originated on the same area; or one may have diverged from the other before it migrated from that area; or even if, when it migrated, it was unchanged, and if in its new home it afterwards split into two species by physiological selection, the newer species would probably prove infertile, not only with its parent type, but also with its grand-parent in any other part of the world.
Seebohm on Isolation.—Seebohm is so strongly influenced by the difficulty from "the swamping effects of free intercrossing," that he is driven by it to adopt Asa Gray's hypothesis of variations as teleological. Indeed, he goes as far as Wagner, for he maintains that in no case can there be divergence or multiplication of species without isolation. He makes the important statement that "the more the geographical distribution of birds is studied, the more doubtful it seems to be that any species of bird has ever been differentiated without the aid of geographical isolation" (Charadriidae, p. 17). If this is true, it makes in favour of physiological selection by showing the paramount importance of the swamping effects of intercrossing, and consequent importance of isolation. But it makes against physiological selection by showing that the geographical form of isolation is sufficient to explain all the cases of specific differentiation in birds. But I must remember that the latter point rests largely on negative inference, and that birds, owing to their highly locomotive habits, are the class of animals where physiological selection is likely to be most handicapped.