With forms which must be ranked as undoubted species, a perfect series exists from those which are absolutely sterile when crossed, to those which are almost or completely fertile. The degrees of sterility do not coincide strictly with the degrees of difference between the parents in external structure or habits of life. Man in many respects may be compared with those animals which have long been domesticated, and a large body of evidence can be advanced in favour of the Pallasian doctrine (14. The 'Variation of Animals and Plants under Domestication,' vol. ii. p. 109. I may here remind the reader that the sterility of species when crossed is not a specially-acquired quality, but, like the incapacity of certain trees to be grafted together, is incidental on other acquired differences. The nature of these differences is unknown, but they relate more especially to the reproductive system, and much less so to external structure or to ordinary differences in constitution. One important element in the sterility of crossed species apparently lies in one or both having been long habituated to fixed conditions; for we know that changed conditions have a special influence on the reproductive system, and we have good reason to believe (as before remarked) that the fluctuating conditions of domestication tend to eliminate that sterility which is so general with species, in a natural state, when crossed. It has elsewhere been shewn by me (ibid. vol. ii. p. 185, and 'Origin of Species,' 5th edit. p. 317), that the sterility of crossed species has not been acquired through natural selection: we can see that when two forms have already been rendered very sterile, it is scarcely possible that their sterility should be augmented by the preservation or survival of the more and more sterile individuals; for, as the sterility increases, fewer and fewer offspring will be produced from which to breed, and at last only single individuals will be produced at the rarest intervals. But there is even a higher grade of sterility than this. Both Gartner and Kolreuter have proved that in genera of plants, including many species, a series can be formed from species which, when crossed, yield fewer and fewer seeds, to species which never produce a single seed, but yet are affected by the pollen of the other species, as shewn by the swelling of the germen. It is here manifestly impossible to select the more sterile individuals, which have already ceased to yield seeds; so that the acme of sterility, when the germen alone is affected, cannot have been gained through selection. This acme, and no doubt the other grades of sterility, are the incidental results of certain unknown differences in the constitution of the reproductive system of the species which are crossed.), that domestication tends to eliminate the sterility which is so general a result of the crossing of species in a state of nature. From these several considerations, it may be justly urged that the perfect fertility of the intercrossed races of man, if established, would not absolutely preclude us from ranking them as distinct species.
Independently of fertility, the characters presented by the offspring from a cross have been thought to indicate whether or not the parent-forms ought to be ranked as species or varieties; but after carefully studying the evidence, I have come to the conclusion that no general rules of this kind can be trusted. The ordinary result of a cross is the production of a blended or intermediate form; but in certain cases some of the offspring take closely after one parent-form, and some after the other. This is especially apt to occur when the parents differ in characters which first appeared as sudden variations or monstrosities. (15. 'The Variation of Animals,' etc., vol. ii. p. 92.) I refer to this point, because Dr. Rohlfs informs me that he has frequently seen in Africa the offspring of negroes crossed with members of other races, either completely black or completely white, or rarely piebald.