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the hinny, which is the offspring of the female ass and stallion. Much stress has been laid by some authors on the supposed fact, that it is only with mongrels that the offspring are not intermediate in character, but closely resemble one of their parents; but this does sometimes occur with hybrids, yet I grant much less frequently than with mongrels. Looking to the cases which I have collected of cross-bred animals closely resembling one parent, the resemblances seem chiefly confined to characters almost monstrous in their nature, and which have suddenly appeared—such as albinism, melanism, deficiency of tail or horns, or additional fingers and toes; and do not relate to characters which have been slowly acquired through selection. A tendency to sudden reversions to the perfect character of either parent would, also, be much more likely to occur with mongrels, which are descended from varieties often suddenly produced and semi-monstrous in character, than with hybrids, which are descended from species slowly and naturally produced. On the whole, I entirely agree with Dr. Prosper Lucas, who, after arranging an enormous body of facts with respect to animals, comes to the conclusion that the laws of resemblance of the child to its parents are the same, whether the two parents differ little or much from each other, namely, in the union of individuals of the same variety, or of different varieties, or of distinct species. Independently of the question of fertility and sterility, in all other respects there seems to be a general and close similarity in the offspring of crossed species, and of crossed varieties. If we look at species as having been specially created, and at varieties as having been produced by secondary laws, this similarity would be an astonishing fact. But it harmonises perfectly with the view that there is no essential distinction between species and varieties.
Summary of Chapter.
First crosses between forms, sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the most careful experimentalists have arrived at diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible to the action of favourable and unfavourable conditions. The degree of sterility does not strictly follow systematic affinity, but is governed by several curious and complex laws. It is generally different, and sometimes widely different in reciprocal crosses between the same two species. It is not always equal in degree in a first cross and in the hybrids produced from this cross.
In the same manner as in grafting trees, the capacity in one species or variety to take on another, is incidental on differences, generally of an unknown nature, in their vegetative systems, so in crossing, the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. There is no more reason to think that species have been specially endowed with various degrees of sterility to prevent their crossing and blending in nature, than to think that trees have been specially endowed with various and somewhat analogous degrees of difficulty in being grafted together in order to prevent their inarching in our forests. The sterility of first crosses and of their hybrid progeny has not been acquired through natural selection. In the case of first crosses it seems to depend on several circumstances; in some instances in chief part on the early death of the embryo. In the case of hybrids, it apparently depends on their whole organisation having been disturbed by being compounded from two distinct forms; the sterility being closely allied to that which so frequently affects pure species, when exposed to new and unnatural conditions of life. He who will explain these latter cases will be able to explain the sterility of hybrids. This view is strongly supported by a parallelism of another kind: namely, that, firstly, slight changes in the conditions of life add to the vigour and fertility of all organic beings; and secondly, that the crossing of forms, which have been exposed to slightly different conditions of life or which have varied, favours the size, vigour, and fertility of their offspring. The facts given on the sterility of the illegitimate unions of dimorphic and trimorphic plants and of their illegitimate progeny, perhaps render it probable that some unknown bond in all cases connects the degree of fertility of first unions with that of their offspring. The consideration of these facts on dimorphism, as well as of the results of reciprocal crosses, clearly leads to the conclusion that the primary cause of the sterility of crossed species is confined to differences in their sexual elements. But why, in the case of distinct species, the sexual elements should so generally have become more or less modified, leading to their mutual infertility, we do not know; but it seems to stand in some close relation to species having been exposed for long periods of time to nearly uniform conditions of life. It is not surprising that the difficulty in crossing any two species, and the sterility of their hybrid offspring, should in most cases correspond, even if due to distinct causes: for both depend on the amount of difference between the species which are crossed. Nor is it surprising that the facility of effecting a first cross, and the fertility of the hybrids thus produced, 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 subjected to experiment; for systematic affinity includes resemblances of all kinds. 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, as is so often stated, invariably fertile. Nor is this almost universal and perfect fertility surprising, when it is remembered 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 that they have not been long exposed to uniform conditions of life. It should also be especially kept in mind, that long-continued domestication tends to eliminate sterility, and is therefore little likely to induce this same quality. Independently of the question of fertility, in all other respects there is the closest general resemblance between hybrids and mongrels, in their variability, in their power of absorbing each other by repeated crosses, and in their inheritance of characters from both parentforms. Finally, then, although we are as ignorant of the precise cause of the sterility of first crosses and of hybrids as we are why animals and plants removed from their natural conditions become sterile, yet the facts given in this chapter do not seem to me opposed to the belief that species aboriginally existed as varieties.