isinclination to cross seems to have been independently acquired, probably by Natural Selection; and I do not see why it would not have sufficed to have prevented incipient species from blending to have simply increased sexual disinclination to cross.
(Paragraph 11.) I demur to a certain extent to amount of sterility and structural dissimilarity necessarily going together, except indirectly and by no means strictly. Look at vars. of pigeons, fowls, and cabbages.
I overlooked the advantage of the half-sterility of reciprocal crosses; yet, perhaps from novelty, I do not feel inclined to admit probability of Natural Selection having done its work so queerly.
I will not discuss the second case of utter sterility, but your assumptions in Paragraph 13 seem to me much too complicated. I cannot believe so universal an attribute as utter sterility between remote species was acquired in so complex a manner. I do not agree with your rejoinder on grafting: I fully admit that it is not so closely restricted as crossing, but this does not seem to me to weaken the case as one of analogy. The incapacity of grafting is likewise an invariable attribute of plants sufficiently remote from each other, and sometimes of plants pretty closely allied.
The difficulty of increasing the sterility through Natural Selection of two already sterile species seems to me best brought home by considering an actual case. The cowslip and primrose are moderately sterile, yet occasionally produce hybrids. Now these hybrids, two or three or a dozen in a whole parish, occupy ground which might have been occupied by either pure species, and no doubt the latter suffer to this small extent. But can you conceive that any individual plants of the primrose and cowslip which happened to be mutually rather more sterile (i.e. which, when crossed, yielded a few less seed) than usual, would profit to such a degree as to increase in number to the ultimate exclusion of the present primrose and cowslip? I cannot.
My son, I am sorry to say, cannot see the full force of your rejoinder in regard to second head of continually augmented sterility. You speak in this rejoinder, and in Paragraph 5, of all the individuals becoming in some slight degree sterile in certain districts: if you were to admit that by continued exposure to these same conditions the sterility would inevitably increase, there would be no need of Natural Selection. But I suspect that the sterility is not caused so much by any particular conditions as by long habituation to conditions of any kind. To speak according to pangenesis, the gemmules of hybrids are not injured, for hybrids propagate freely by buds; but their reproductive organs are somehow affected, so that they cannot accumulate the proper gemmules, in nearly the same manner as the reproductive organs of a pure species become affected when exposed to unnatural conditions.
This is a very ill-expressed and ill-written letter. Do not answer it, unless the spirit urges you. Life is too short for so long a discussion. We shall, I greatly fear, never agree.
LETTER 214. A.R. WALLACE TO CHARLES DARWIN. Hurstpierpoint, [April?] 8th, 1868.
I am sorry you should have given yourself the trouble to answer my ideas on sterility. If you are not convinced, I have little doubt but that I am wrong; and, in fact, I was only half convinced by my own arguments, and I now think there is about an even chance that Natural Selection may or may not be able to accumulate sterility. If my first proposition is modified to the existence of a species and a variety in the same area, it will do just as well for my argument. Such certainly do exist. They are fertile together, and yet each maintains itself tolerably distinct. How can this be, if there is no disinclination to crossing?
My belief certainly is that number of offspring is not so important an element in keeping up population of a species as supply of food and other favourable conditions; because the numbers of a species constantly vary greatly in different parts of its own area, whereas the average number of offspring is not a very variable element.