This greater variability of mongrels than of hybrids does not seem to me at all surprising. For the parents of mongrels are varieties, and mostly domestic varieties (very few experiments having been tried on natural varieties), and this implies in most cases that there has been recent variability; and therefore we might expect that such variability would often continue and be superadded to that arising from the mere act of crossing. The slight degree of variability in hybrids from the first cross or in the first generation, in contrast with their extreme variability in the succeeding generations, is a curious fact and deserves attention. For it bears on and corroborates the view which I have taken on the cause of ordinary variability; namely, that it is due to the reproductive system being eminently sensitive to any change in the conditions of life, being thus often rendered either impotent or at least incapable of its proper function of producing offspring identical with the parent-form. Now hybrids in the first generation are descended from species (excluding those long cultivated) which have not had their reproductive systems in any way affected, and they are not variable; but hybrids themselves have their reproductive systems seriously affected, and their descendants are highly variable.

But to return to our comparison of mongrels and hybrids Gärtner states that mongrels are more liable than hybrids to revert to either parent-form; but this, if it be true, is certainly only a difference in degree. Moreover, Gärtner expressly states that hybrids from longcultivated plants are more subject to reversion than hybrids from species in their natural state; and this probably explains the singular difference in the results arrived at by different observers: thus, Max Wichura doubts whether hybrids ever revert to their parent

forms, and he experimented on uncultivated species of willows; whilst Naudin, on the other hand, insists in the strongest terms on the almost universal tendency to reversion in hybrids, and he experimented chiefly on cultivated plants. Gärtner further states that when any two species, although most closely allied to each other, are crossed with a third species, the hybrids are widely different from each other; whereas, if two very distinct varieties of one species are crossed with another species, the hybrids do not differ much. But this conclusion, as far as I can make out, is founded on a single experiment; and seems directly opposed to the results of several experiments made by Kölreuter.

Such alone are the unimportant differences which Gärtner is able to point out between hybrid and mongrel plants. On the other hand, the degrees and kinds of resemblance in mongrels and in hybrids to their respective parents, more especially in hybrids produced from nearly related species, follow according to Gärtner the same laws. When two species are crossed, one has sometimes a prepotent power of impressing its likeness on the hybrid; and so I believe it to be with varieties of plants. With animals one variety certainly often has this prepotent power over another variety. Hybrid plants produced from a reciprocal cross, generally resemble each other closely; and so it is with mongrels from a reciprocal cross. Both hybrids and mongrels can be reduced to either pure parent-form, by repeated crosses in successive generations with either parent.

These several remarks are apparently applicable to animals; but the subject is here much complicated, partly owing to the existence of secondary sexual characters; but more especially owing to prepotency in transmitting likeness running more strongly in one

sex than in the other, both when one species is crossed with another, and when one variety is crossed with another variety. For instance, I think those authors are right, who maintain that the ass has a prepotent power over the horse, so that both the mule and the hinny more resemble the ass than the horse; but that the prepotency runs more strongly in the male-ass than in the female, so that the mule, which is the offspring of the male-ass and mare, is more like an ass, than is 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 with them 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. Consequently, sudden reversions to the perfect character of either parent would be 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.

Laying aside 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 two most careful experimentalists who haye ever lived, have come to diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible 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 of one species or variety to take on another, is incidental on generally unknown differences 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 them 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 them becoming inarched in our forests.

The sterility of first crosses and of their hybrid progeny has not, as far as we can judge, been increased through natural selection so as to attain that high degree which is universal with species when rendered widely distinct. With first crosses between pure species, in which the reproductive system is in a perfect condition, the sterility seems to depend on several circumstances; in some cases largely on the early death of the embryo, but this apparently depends on some imperfection in the original act of impregnation. With hybrids, in which the reproductive system is in an imperfect condition, and in which this system as well as the whole organisation has been disturbed by being compounded from two distinct forms, the sterility apparently is closely allied to that which so frequently affects pure species, when exposed to unnatural conditions of life. This view is supported by a parallelism of another kind: namely, that the crossing of forms only slightly differentiated favours the vigour and fertility of their offspring; and that slight changes in the conditions of life apparently add to the vigour and fertility of all organic beings. But the facts given on the sterility of the illegitimate unions of dimorphic and trimorphic plants and of their illegitimate progeny, render it probable that there is some unknown bond connecting in all cases the degree of fertility of first unions with that of their offspring. The consideration of these facts on dimorphism, as well as the results of reciprocal crosses, drive us to conclude that in all cases the primary cause