Origin of Species Variorum

It is, no doubt, extremely difficult even to conjecture by what gradations many structures have been perfected, more especially amongst broken and failing groups of organic beings, which have suffered much extinction; but we see so many strange gradations in nature, ... that we ought to be extremely cautious in saying that any organ or instinct, or any whole being, could not have arrived at its present state by many graduated steps. There are, it must be admitted, cases of special difficulty on the theory of natural selection; and one of the most curious of these is the existence of two or three defined castes of workers or sterile females in the same community of ants; but I have attempted to show how these difficulties can be mastered.

With respect to the almost universal sterility of species when first crossed, which forms so remarkable a contrast with the almost universal fertility of varieties when crossed, I must refer the reader to the recapitulation of the facts given at the end of the eighth chapter, which seem to me conclusively to show that this sterility is no more a special endowment than is the incapacity of two trees to be grafted together; but that it is incidental on constitutional differences in the reproductive systems of the intercrossed species. We see the truth of this conclusion in the vast difference in the result, when the same two species are crossed reciprocally; that is, when one species is first used as the father and then as the mother. ↑

Although the fertility of varieties when intercrossed and of their mongrel offspring has been asserted by so many authors to be universal, this cannot be considered correct after the facts given on the authority of Gärtner and Kölreuter. Nor is the very general fertility of varieties, when crossed, surprising, when we remember that it is not likely that either their constitutions or their reproductive systems should have been profoundly modified. Moreover, most of the varieties which have been experimented on have been produced under domestication; and as domestication (I do not mean mere confinement) apparently tends to eliminate sterility, we