In all these respects, and in others which might be added, the forms of the same undoubted species, when illegitimately united, behave in exactly the same manner as do two distinct species when crossed. This led me carefully to observe during four years many seedlings, raised from several illegitimate unions. The chief result is that these illegitimate plants, as they may be called, are not fully fertile. It is possible to raise from dimorphic species, both long-styled and short-styled illegitimate plants, and from trimorphic plants all three illegitimate forms. These can then be properly united in a legitimate manner. When this is done, there is no apparent reason why they should not yield as many seeds as did their parents when legitimately fertilised. But such is not the case. They are all infertile, in various degrees; some being so utterly and incurably sterile that they did not yield during four seasons a single seed or even seed-capsule. The sterility of these illegitimate plants, when united with each other in a legitimate manner, may be strictly compared with that of hybrids when crossed inter se. If, on the other hand, a hybrid is crossed with either pure parent-species, the sterility is usually much lessened: and so it is when an illegitimate plant is fertilised by a legitimate plant. In the same manner as the sterility of hybrids does not always run parallel with the difficulty of making the first cross between the two parent-species, so that sterility of certain illegitimate plants was unusually great, while the sterility of the union from which they were derived was by no means great. With hybrids raised from the same seed- capsule the degree of sterility is innately variable, so it is in a marked manner with illegitimate plants. Lastly, many hybrids are profuse and persistent flowerers, while other and more sterile hybrids produce few flowers, and are weak, miserable dwarfs; exactly similar cases occur with the illegitimate offspring of various dimorphic and trimorphic plants.

Altogether there is the closest identity in character and behaviour between illegitimate plants and hybrids. It is hardly an exaggeration to maintain that illegitimate plants are hybrids, produced within the limits of the same species by the improper union of certain forms, while ordinary hybrids are produced from an improper union between so-called distinct species. We have also already seen that there is the closest similarity in all respects between first illegitimate unions and first crosses between distinct species. This will perhaps be made more fully apparent by an illustration; we may suppose that a botanist found two well-marked varieties (and such occur) of the long-styled form of the trimorphic Lythrum salicaria, and that he determined to try by crossing whether they were specifically distinct. He would find that they yielded only about one-fifth of the proper number of seed, and that they behaved in all the other above specified respects as if they had been two distinct species. But to make the case sure, he would raise plants from his supposed hybridised seed, and he would find that the seedlings were miserably dwarfed and utterly sterile, and that they behaved in all other respects like ordinary hybrids. He might then maintain that he had actually proved, in accordance with the common view, that his two varieties were as good and as distinct species as any in the world; but he would be completely mistaken.

The facts now given on dimorphic and trimorphic plants are important, because they show us, first, that the physiological test of lessened fertility, both in first crosses and in hybrids, is no safe criterion of specific distinction; secondly, because we may conclude that there is some unknown bond which connects the infertility of illegitimate unions with that of their illegitimate offspring, and we are led to extend the same view to first crosses and hybrids; thirdly, because we find, and this seems to me of especial importance, that two or three forms of the same species may exist and may differ in no respect whatever, either in structure or in constitution, relatively to external conditions, and yet be sterile when united in certain ways. For we must remember that it is the union of the sexual elements of individuals of the same form, for instance, of two long- styled forms, which results in sterility; while it is the union of the sexual elements proper to two distinct forms which is fertile. Hence the case appears at first sight exactly the reverse of what occurs, in the ordinary unions of the individuals of the same species and with crosses between distinct species. It is, however, doubtful whether this is really so; but I will not enlarge on this obscure subject.

We may, however, infer as probable from the consideration of dimorphic and trimorphic plants, that the sterility of distinct species when crossed and of their hybrid progeny, depends exclusively on the nature of their sexual elements, and not on any difference in their structure or general constitution. We are also led to this same conclusion by considering reciprocal crosses, in which the male of one species cannot be united, or can be united with great difficulty, with the female of a second species, while the converse cross can be effected with perfect facility. That excellent observer, Gartner, likewise concluded that species when crossed are sterile owing to differences confined to their reproductive systems.

FERTILITY OF VARIETIES WHEN CROSSED, AND OF THEIR MONGREL OFFSPRING, NOT UNIVERSAL.

It may be urged as an overwhelming argument that there must be some essential distinction between species and varieties inasmuch as the latter, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring. With some exceptions, presently to be given, I fully admit that this is the rule. But the subject is surrounded by difficulties, for, looking to varieties produced under nature, if two forms hitherto reputed to be varieties be found in any degree sterile together, they are at once ranked by most naturalists as species. For instance, the blue and red pimpernel, which are considered by most botanists as varieties, are said by Gartner to be quite sterile when crossed, and he consequently ranks them as undoubted species. If we thus argue in a circle, the fertility of all varieties produced under nature will assuredly have to be granted.

If we turn to varieties, produced, or supposed to have been produced, under domestication, we are still involved in some doubt. For when it is stated, for instance, that certain South American indigenous domestic dogs do not readily unite with European dogs, the explanation which will occur to everyone, and probably the true one, is that they are descended from aboriginally distinct species. Nevertheless the perfect fertility of so many domestic races, differing widely from each other in appearance, for instance, those of the pigeon, or of the cabbage, is a remarkable fact; more especially when we reflect how many species there are, which, though resembling each other most closely, are utterly sterile when intercrossed. Several considerations, however, render the fertility of domestic varieties less remarkable. In the first place, it may be observed that the amount of external difference between two species is no sure guide to their degree of mutual sterility, so that similar differences in the case of varieties would be no sure guide. It is certain that with species the cause lies exclusively in differences in their sexual constitution. Now the varying conditions to which domesticated animals and cultivated plants have been subjected, have had so little tendency towards modifying the reproductive system in a manner leading to mutual sterility, that we have good grounds for admitting the directly opposite doctrine of Pallas, namely, that such conditions generally eliminate this tendency; so that the domesticated descendants of species, which in their natural state probably would have been in some degree sterile when crossed, become perfectly fertile together. With plants, so far is cultivation from giving a tendency towards sterility between distinct species, that in several well- authenticated cases already alluded to, certain plants have been affected in an opposite manner, for they have become self-impotent, while still retaining the capacity of fertilising, and being fertilised by, other species. If the Pallasian doctrine of the elimination of sterility through long-continued domestication be admitted, and it can hardly be rejected, it becomes in the highest degree improbable that similar conditions long- continued should likewise induce this tendency; though in certain cases, with species having a peculiar constitution, sterility might occasionally be thus caused. Thus, as I believe, we can understand why, with domesticated animals, varieties have not been produced which are mutually sterile; and why with plants only a few such cases, immediately to be given, have been observed.

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