Dean Herbert, who carried on experiments with great care and skill for many years, found numerous cases of hybrids which were perfectly fertile inter se. Crinum capense, fertilised by three other species—C. pedunculatum, C. canaliculatum, or C. defixum—all very distinct from it, produced perfectly fertile hybrids; while other species less different in appearance were quite sterile with the same C. capense.
All the species of the genus Hippeastrum produce hybrid offspring which are invariably fertile. Lobelia syphylitica and L. fulgens, two very distinct species, have produced a hybrid which has been named Lobelia speciosa, and which reproduces itself abundantly. Many of the beautiful pelargoniums of our greenhouses are hybrids, such as P. ignescens from a cross between P. citrinodorum and P. fulgidum, which is quite fertile, and has become the parent of innumerable varieties of beautiful plants. All the varied species of Calceolaria, however different in appearance, intermix with the greatest readiness, and the hybrids are all more or less fertile. But the most remarkable case is that of two species of Petunia, of which Dean Herbert says: "It is very remarkable that, although there is a great difference in the form of the flower, especially of the tube, of P. nyctanigenaeflora and P. phoenicea the mules between them are not only fertile, but I have found them seed much more freely with me than either parent.... From a pod of the above-mentioned mule, to which no pollen but its own had access, I had a large batch of seedlings in which there was no variability or difference from itself; and it is evident that the mule planted by itself, in a congenial climate, would reproduce itself as a species; at least as much deserving to be so considered as the various Calceolarias of different districts of South America."[56]
Darwin was informed by Mr. C. Noble that he raises stocks for grafting from a hybrid between Rhododendron ponticum and R. catawbiense, and that this hybrid seeds as freely as it is possible to imagine. He adds that horticulturists raise large beds of the same hybrid, and such alone are fairly treated; for, by insect agency, the several individuals are freely crossed with each other, and the injurious influence of close interbreeding is thus prevented. Had hybrids, when fairly treated, always gone on decreasing in fertility in each successive generation, as Gartner believed to be the case, the fact would have been notorious to nurserymen.[57]
Cases of Sterility of Mongrels.
The reverse phenomenon to the fertility of hybrids, the sterility of mongrels or of the crosses between varieties of the same species, is a comparatively rare one, yet some undoubted cases have occurred. Gartner, who believed in the absolute distinctness of species and varieties, had two varieties of maize—one dwarf with yellow seeds, the other taller with red seeds; yet they never naturally crossed, and, when fertilised artificially, only a single head produced any seeds, and this one only five grains. Yet these few seeds were fertile; so that in this case the first cross was almost sterile, though the hybrid when at length produced was fertile. In like manner, dissimilarly coloured varieties of Verbascum or mullein have been found by two distinct observers to be comparatively infertile. The two pimpernels (Anagallis arvensis and A. coerulea), classed by most botanists as varieties of one species, have been found, after repeated trials, to be perfectly sterile when crossed.
No cases of this kind are recorded among animals; but this is not to be wondered at, when we consider how very few experiments have been made with natural varieties; while there is good reason for believing that domestic varieties are exceptionally fertile, partly because one of the conditions of domestication was fertility under changed conditions, and also because long continued domestication is believed to have the effect of increasing fertility and eliminating whatever sterility may exist. This is shown by the fact that, in many cases, domestic animals are descended from two or more distinct species. This is almost certainly the case with the dog, and probably with the hog, the ox, and the sheep; yet the various breeds are now all perfectly fertile, although we have every reason to suppose that there would be some degree of infertility if the several aboriginal species were crossed together for the first time.
Parallelism between Crossing and Change of Conditions.
In the whole series of these phenomena, from the beneficial effects of the crossing of different stocks and the evil effects of close interbreeding, up to the partial or complete sterility induced by crosses between species belonging to different genera, we have, as Mr. Darwin points out, a curious parallelism with the effects produced by change of physical conditions. It is well known that slight changes in the conditions of life are beneficial to all living things. Plants, if constantly grown in one soil and locality from their own seeds, are greatly benefited by the importation of seed from some other locality. The same thing happens with animals; and the benefit we ourselves experience from "change of air" is an illustration of the same phenomenon. But the amount of the change which is beneficial has its limits, and then a greater amount is injurious. A change to a climate a few degrees warmer or colder may be good, while a change to the tropics or to the arctic regions might be injurious.
Thus we see that, both slight changes of conditions and a slight amount of crossing, are beneficial; while extreme changes, and crosses between individuals too far removed in structure or constitution, are injurious. And there is not only a parallelism but an actual connection between the two classes of facts, for, as we have already shown, many species of animals and plants are rendered infertile, or altogether sterile, by the change from their natural conditions which occurs in confinement or in cultivation; while, on the other hand, the increased vigour or fertility which is invariably produced by a judicious cross may be also effected by a judicious change of climate and surroundings. We shall see in a subsequent chapter, that this interchangeability of the beneficial effects of crossing and of new conditions, serves to explain some very puzzling phenomena in the forms and economy of flowers.
Remarks on the Facts of Hybridity.