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 organisation 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 their 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 the 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. Finally, then, the facts briefly given in this chapter do not seem to me opposed to, but even rather to support the view, that there is no fundamental distinction between species and varieties.
CHAPTER IX.
On the Imperfection of the Geological Record.
On the absence of intermediate varieties at the present day—On the nature of extinct intermediate varieties; on their number—On the vast lapse of time, as inferred from the rate of deposition and of denudation—On the poorness of our palæontological collections—On the intermittence of geological formations—On the absence of intermediate varieties in any one formation—On the sudden appearance of groups of species—On their sudden appearance in the lowest known fossiliferous strata.
In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. Most of them have now been discussed. One, namely the distinctness of specific forms, and their not being blended together by innumerable transitional links, is a very obvious difficulty. I assigned reasons why such links do not commonly occur at the present day, under the circumstances apparently most favourable for their presence, namely on an extensive and continuous area with graduated physical conditions. I endeavoured to show, that the life of each species depends in a more important manner on the presence of other already defined organic forms, than on climate; and, therefore, that the really governing conditions of life do not graduate away quite insensibly like heat or moisture. I endeavoured, also, to show that intermediate varieties, from existing in lesser numbers than the forms which they connect, will generally be beaten out and exterminated during the course of further modification and improvement. The main cause, however, of innumerable intermediate links not now occurring everywhere throughout nature
depends on the very process of natural selection, through which new varieties continually take the places of and exterminate their parent-forms. But just in proportion as this process of extermination has acted on an enormous scale, so must the number of intermediate varieties, which have formerly existed on the earth, be truly enormous. Why then is not every geological formation and every stratum full of such intermediate links? Geology assuredly does not reveal any such finely graduated organic chain; and this, perhaps, is the most obvious and gravest objection which can be urged against my theory. The explanation lies, as I believe, in the extreme imperfection of the geological record.