To avoid complication, it must be granted, that between As and Bs no cross-unions take place, while between Af and Bf cross-unions are as frequent as direct unions, though much less fertile. We must also leave out of consideration crosses between As and Af, Bs and Bf, with their various approaches to sterility, as I believe they will not affect the final result, although they will greatly complicate the problem.
14. In the first generation there will result: 1st, The pure progeny of As and Bs; 2nd, The pure progeny of Af and of Bf; and 3rd, The hybrid progeny of Af, Bf.
15. Supposing that, in ordinary years, the increased constitutional vigour of the hybrids exactly counterbalances their imperfect adaptations to conditions, there will be in the second generation, besides these three classes, hybrids of the second degree between the first hybrids and Af and Bf respectively. In succeeding generations there will be hybrids of all degrees, varying between the first hybrids and the almost pure types of Af and Bf.
16. Now, if at first the number of individuals of As, Bs, Af and Bf were equal, and year after year the total number continues stationary, I think it can be proved that, while half will be the pure progeny of As and Bs, the other half will become more and more hybridised, until the whole will be hybrids of various degrees.
17. Now, this hybrid and somewhat intermediate race cannot be so well adapted to the conditions of life as the two pure species, which have been formed by the minute adaptation to conditions through Natural Selection; therefore, in a severe struggle for existence, the hybrids must succumb, especially as, by hypothesis, their fertility would not be so great as that of the two pure species.
18. If we were to take into consideration the unions of As with Af and Bs with Bf, the results would become very complicated, but it must still lead to there being a number of pure forms entirely derived from As and Bs, and of hybrid forms mainly derived from Af and Bf; and the result of the struggle of these two sets of individuals cannot be doubtful.
19. If these arguments are sound, it follows that sterility may be accumulated and increased, and finally made complete by Natural Selection, whether the sterile varieties originate together in a definite portion of the area occupied by the two species, or occur scattered over the whole area. (211/4. The first part of this discussion should be considered alone, as it is both more simple and more important. I now believe that the utility, and therefore the cause of sterility between species, is during the process of differentiation. When species are fully formed, the occasional occurrence of hybrids is of comparatively small importance, and can never be a danger to the existence of the species. A.R.W. (1899).)
P.S.--In answer to the objection as to the unequal sterility of reciprocal crosses ("Variation, etc." Volume II., page 186) I reply that, as far as it went, the sterility of one cross would be advantageous even if the other cross was fertile: and just as characters now co-ordinated may have been separately accumulated by Natural Selection, so the reciprocal crosses may have become sterile one at a time.
LETTER 212. TO A.R. WALLACE. 4, Chester Place, March 17th, 1868.
(212/1. Mr. Darwin had already written a short note to Mr. Wallace expressing a general dissent from his view.)
I do not feel that I shall grapple with the sterility argument till my return home; I have tried once or twice, and it has made my stomach feel as if it had been placed in a vice. Your paper has driven three of my children half mad--one sat up till 12 o'clock over it. My second son, the mathematician, thinks that you have omitted one almost inevitable deduction which apparently would modify the result. He has written out what he thinks, but I have not tried fully to understand him. I suppose that you do not care enough about the subject to like to see what he has written.
LETTER 212A.