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have seen, the spherical surfaces would wholly disappear, and would all be replaced by plane surfaces ; and the Melipona would make a comb as perfect as that of the hive-bee. Beyond this stage of perfection in architecture, natural selection could not load ; for the comb of the hive-bee, as far as we can see, is absolutely perfect in economising wax.
Thus, as I believe, the most wonderful of all known instincts, that of the hive-bee, can be explained by natural selection having taken advantage of numerous, successive, slight modifications of simpler instincts; natural selection having by slow degreos, more and more perfectly, led the bees to sweep equal spheres at a given distance from each other in a double layer, and to build up and excavate the wax along the planes of intersection. The bees, of course, no more knowing that they swept their spheres at one particular distance from each other, than they know what are the several angles of the hexagonal prisms and of the basal rhombic plates. The motive power of the process of natural selection having been economy of wax; that individual swarm which wasted least honey in the secretion of wax, having succeeded best, and having transmitted by inheritance its newly acquired economical instinct to new swarms, which in their turn will have had the best chance of succeeding in the struggle for existence.
No doubt many instincts of very difficult explanation could be opposed to the theory of natural selection, cases, in which we cannot see how an instinct could possibly have originated ; cases, in which no intermediate gradations are known to exist; cases of instinct of apparently such trifling importance, that they could hardly have been acted on by natural selection ; cases of instincts almost identically the same in animals so remote in the scale of nature, that we cannot account for their similarity by inheritance from a common parent, and must therefore believe that they have been acquired by independent acts of natural selection.
I will not here enter on these several cases, but will confine myself to one special difficulty, which at first appeared to me insuperable, and actually fatal to my whole theory. I allude to the neuters or sterile females in insect-communities : for these neuters often differ widely in instinct and in structure from both the males and fertile females, and yet, from being sterile, they cannot propagate their kind.
The subject well deserves to be discussed at great length, but I will here take only a single case, that of working or sterile ants. How the workers have been rendered sterile is a difficulty ; but not much greater than that of any other striking modification of structure ; for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile; and if such insects had been social, and it had been profitable to the community that a number should have been annually born capable of work, but incapable of procreation, I can see no very great difficulty in this being effected by natural selection. But I must pass over this preliminary difficulty. The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure, as in the shape of the thorax and in being destitute of wings and sometimes of eyes, and in instinct. As far as instinct alone is concerned, the prodigious difference in this respect between the workers and the perfect females, would have been far better exemplified by the hive-bee. If a working ant or other neuter insect had been an animal in tho ordinary state, I should have unhesitatingly, assumed that all its characters had been slowly acquired through natural selection ; namely, by an individual having been born with some slight profitable modification of structure, this being inherited by its offspring, which again varied and were again selected, and so onwards. But with the working ant we have an insect differing greatly from its parents, yet absolutely sterile; so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny.
It may well be asked how is it possible to reconcile this case with the theory of natural selection?
First, let it be remembered that we have innumerable instances, both in our domestic productions and in those in a state of nature, of all sorts of differences of structure which have become correlated to certain ages, and to either sex. We have differences correlated not only to one sex, but to that short period alone when the reproductive system is active, as in the nuptial plumage of many birds, and in the hooked jaws of the male salmon. We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the malo sex; for oxen of certain breeds have longer horns than in other breeds, in comparison with the horns of the bulls or cows of these same breeds. Hence I can see no real difficulty in any character having become correlated with the sterile condition of certain members of insect - communities: the difficulty lies in understanding how such correlated modifications of structure could have been slowly accumulated by natural selection.
This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end. Thus, a well - flavoured vegetable is cooked, and the individual is destroyed; but the horticulturist sows seeds of the same stock, and confidently expects to get nearly the same variety : breeders of cattle wish the flesh and fat to be well marbled together; the animal has been slaughtered, but the breeder goes with confidence to the same family. I have such faith in the powers of selection, that I do not doubt that a breed of cattle, always yielding oxen with extraordinarily long horns, could be slowly formed by carefully watching which individual bulls and cows, when matched, produced oxen with the longest horns; and yet no one ox could ever have propagated its kind. Thus I believe it bas been
with social insects : a slight modification of structure, or instinct, correlated with the sterile condition of certain members of the community, has been advantageous to the community: consequently the fertile males and females of the same community flourished, and transmitted to their fertile offspring a tendency to produce sterile members having the same modification. And I believe that this process has been repeated, until that prodigious amount of difference between the fertile and sterile females of the same species has been produced, which we see in many social insects.
But we have not as yet touched on the climax of the difficulty ; namely, the fact that the neuters of several ants differ, not only from the fertile females and
males, but from each other, sometimes to an almost incredible degree, and are thus divided into two or even three castes. The castes, moreover, do not generally graduate into each other, but are perfectly well defined ; being as distinct from each other, as are any two species of the same genus, or rather as any two genera of the same family. Thus in Eciton, there are working and soldier neuters, with jaws and instincts extraordinarily different: in Cryptocerus, the workers of one caste alone carry a wonderful sort of shield on their heads, the use of which is quite unknown : in the Mexican Myrmecocystus, the workers of one caste never leave the nest; they are fed by the workers of another caste, and they have an enormously developed abdomen which secretes a sort of honey, supplying the place of that excreted by the aphides, or the domesti: cattle as they may be called, which our European ants guard or imprison.
It will indeed be thought that I have an overweening confidence in the principle of natural selection, when I do not admit that such wonderful and well-established facts at once annihilate my theory. In the simpler case of neuter insects all of one caste or of the same kind, which have been rendered by natural selection, as I believe to be quite possible, different from the fertile males and females, —in this case, we may safely
conclude from the analogy of ordinary variations, that each successive, slight, profitable modification did not probably at first appear in all the individual neuters in the same nest, but in a few alone; and that by the long-continued selection of the fertile parents which produced most neuters with the profitable modification, all the neuters ultimately came to have the desired character. On this view we ought occasionally to find nouter-insects of the same species, in the same nest, presenting gradations of structure; and this we do find, even often, considering how few neuter-insects out of Europe have been carefully examined. Mr. F. Smith has shown how surprisingly the neuters of severa) British ants differ from each other in size and some times in colour; and that the extreme forms can sometimes be perfectly linked together by individuals taken out of the same nest: I have myself compared perfect gradations of this kind. It often happens that the larger or the smaller sized workers are the most numerous; or that both large and small are numerous, with those of an intermediate size scanty in numbers. Formica flava has, larger and smaller workers, with some of intermediate size; and, in this species, as Mr. F. Smith has observed, the larger workers have simple eyes (ocelli), which though small can be plainly distinguished, whereas the smaller workers have their ocelli rudimentary. Having carefully dissected several specimens of these workers, I can affirm that the eyes are far more rudimentary in the smaller workers than can be accounted for merely by their proportionally lesser size; and I fully believe, though I dare not assert so positively, that the workers of intermediate size have their ocelli in an exactly intermediate condition. So that we here have two bodies of sterile workers in the same nest, differing not only in size, but in their organs of vision, yet connected by some few members in an intermediate condition. digress by adding, that if the smaller workers had been tho most useful to the community, and those males and females had been continually selected, which produced