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a hybrid from Calceolaria integrifolia and plantaginea, species most widely dissimilar in general habit, “ reproduces itself as perfectly as if it had been a natural species from the mountains of Chili.” I have taken some pains to ascertain the degree of fertility of some of the complex crosses of Rhododendrons, and I am assured that many of them are perfectly fertile. Mr. C. Noble, for instance, informs me that he raises stocks for grafting from a hybrid between Rhod. ponticum and catawbiense, and that this hybrid “seeds as freely as it is possible to imagine.” Had hybrids when fairly treated, always gone on decreasing in fertility in each successive generation, as Gärtner believed to be the case, the fact would have been notorious to nurserymen. Horticulturists raise large beds of the same hybrid, and such alone are fairly treated, for by insect agency the several individuals are allowed to cross freely with each other, and the injurious influence of close interbreeding is thus prevented. Any one may readily convince himself of the efficiency of insect-agency by examining the flowers of the more sterile kinds of hybrid Rhododendrons, which produce no pollen, for he will find on their stigmas plenty of pollen brought from other flowers. In regard to animals, much fewer experiments have been carefully tried than with plants. If our systematic arrangements can be trusted, that is, if the genera of animals are as distinct from each other as are the genera of plants, then we may infer that animals more widely distinct in the scale of nature can be crossed more easily than in the case of plants; but the hybrids themselves are, I think, more sterile. It should, however, be borne in mind that, owing to few animals breeding freely under confinement, few experiments have been fairly tried: for instance, the canary-bird has been crossed with nine distinct species of finches, but, as not one of these breeds freely in confinement, we have no right to expect that the first crosses between them and the canary, or that their hybrids, should be perfectly fertile. Again, with respect to the fertility in successive generations of the more fertile hybrid animals, I hardly know of an instance in which two families of the same hybrid have been raised at the same time from different parents, so as to avoid the ill effects of close interbreeding. On the contrary, brothers and sisters have usually been crossed in each successive generation, in opposition to the constantly repeated admonition of every breeder. And in this case, it is not at all surprising that the inherent sterility in the hybrids should have gone on increasling. Although I know of hardly any thoroughly wellauthenticated cases of perfectly fertile hybrid animals, I have reason to believe that the hybrids from Cervulus vaginalis and Reevesii, and from Phasianus colchicus with P. torquatus, are perfectly fertile. M. Quatrefages states that the hybrids from two moths (Bombyx cynthia and arrindia) were proved in Paris to be fertile inter se for eight generations. It has lately been asserted that two such distinct species as the hare and rabbit, when they can be got to breed together, produce offspring, which are highly fertile when crossed with one of the parent-species. The hybrids from the common and Chinese geese (A. cygnoides), species which are so different that they are generally ranked in distinct genera, have often bred in this country with either pure parent, and in one single instance they have bred inter se. This was effected by Mr. Eyton, who raised two hybrids from the same parents, but from different hatches; and from these two birds he raised no less than eight hybrids (grandchildren of the pure geese) from one nest. In India, however, these cross-bred geese must be far more fertile; for I am assured by two eminently capable judges, namely Mr. Blyth and Capt. Hutton, that whole flocks of these crossed geese are kept in various parts of the country; and as they are kept for profit, where neither pure parent-species, exists, they must certainly be highly or perfectly fertile. With our domesticated animals, the various races when crossed together are quite fertile; yet in many cases they are descended from two or more wild species. From this fact we must conclude either that the aboriginal parent-species at first produced perfectly fertile hybrids, or that the hybrids subsequently reared under domestication became quite fertile. This latter alternative, which was first propounded by Pallas, seems by far the most probable, and can, indeed, hardly be doubted. It is, for instance, almost certain that our dogs are descended from several wild stocks; yet, with perhaps the exception of certain indigenous domestic dogs of South America, all are quite fertile together; but analogy makes me greatly doubt, whether the several aboriginal species would at first have freely bred together and have produced quite fertile hybrids. So again I have lately acquired decisive evidence that the crossed offspring from the Indian humped and common cattle are inter se perfectly fertile; and from the observations by Rütimeyer on their important osteological differences, as well as from those by Mr. Blyth on their differences in habits, voice, constitution, &c., these two forms must be regarded as good and distinct species. The same remarks may be extended to the two chief races of the pig. We must, therefore, either give up the belief of the universal sterility of species when crossed; or we must look at this sterility in animals, not as an indelible characteristic, but as one capable of being removed by domestication. Finally, considering all the ascertained facts on the intercrossing of plants and animals, it may be concluded that some degree of sterility, both in first crosses and in hybrids, is an extremely general result; but that it cannot, under our present state of knowledge, be considered as absolutely universal.
Laws governing the Sterility of first Crosses and of Hybrids.
We will now consider a little more in detail the laws governing the sterility of first crosses and of hybrids. Our chief object will be to see whether or not these laws indicate that species have been specially endowed with this quality, in order to prevent their crossing and blending together in utter confusion. The following conclusions are drawn up chiefly from Gärtner's admirable work on the hybridisation of plants. I have taken much pains to ascertain how far they apply to animals, and, considering how scanty our knowledge is in regard to hybrid animals, I have been surprised to find how generally the same rules apply to both kingdoms.
It has been already remarked, that the degree of fertility, both of first crosses and of hybrids, graduates from zero to perfect fertility. It is surprising in how many curious ways this gradation can be shown; but only the barest outline of the facts can here be given. When pollen from a plant of one family is placed on the stigma of a plant of a distinct family, it exerts no more influence than so much inorganic dust. From this absolute zero of fertility, the pollen of different species applied to the stigma of some one species of the same genus, yields a perfect gradation in the number of seeds produced, up to nearly complete or even quite complete fertility; and, as we have seen, in certain abnormal cases, even to an excess of fertility, beyond that which the plant's own pollen produces. So in hybrids themselves, there are some which never have produced, and probably never would produce, even with the pollen of the pure parents, a single fertile seed: but in some of these cases a first trace of fertility may be detected, by the pollen of one of the pure parent-species causing the flower of the hybrid to wither earlier than it otherwise would have done; and the early withering of the flower is well known to be a sign of incipient fertilisation. From this extreme degree of sterility we have self-fertilised hybrids producing a greater and greater number of seeds up to perfect fertility. The hybrids raised from two species which are very difficult to cross, and which rarely produce any offspring, are generally very sterile; but the parallelism between the difficulty of making a first cross, and the sterility of the hybrids thus produced—two classes of facts which are generally confounded together—is by no means strict. There are many cases, in which two pure species, as in the genus Verbascum, can be united with unusual facility, and produce numerous hybrid