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07 - Instinct 07-10 - Neuter or sterile insects 10 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.

thorax
thorax


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.

Hive Bee
Hive Bee


If a working ant or other neuter insect had been an animal in the 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.

plumage
plumage

salmon
salmon

Peacock Female
Peacock Female


We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the male 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.

cattle
cattle


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 has 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 domestic cattle as they may be called, which our European ants guard or imprison.

eciton (Army Ant)
eciton (Army Ant)

cryptocerus
cryptocerus

myrmecocystus
myrmecocystus


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 neuter-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 several British ants differ from each other in size and sometimes 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.

I may digress by adding, that if the smaller workers had been the most useful to the community, and those males and females had been continually selected, which produced more and more of the smaller workers, until all the workers had come to be in this condition; we should then have had a species of ant with neuters very nearly in the same condition with those of Myrmica.

Myrmica Worker
Myrmica Worker


For the workers of Myrmica have not even rudiments of ocelli, though the male and female ants of this genus have well-developed ocelli.

ocelli
ocelli


I may give one other case: so confidently did I expect to find gradations in important points of structure between the different castes of neuters in the same species, that I gladly availed myself of Mr F. Smith's offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa.

Driver Ant
Driver Ant


The reader will perhaps best appreciate the amount of difference in these workers, by my giving not the actual measurements, but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house of whom many were five feet four inches high, and many sixteen feet high; but we must suppose that the larger workmen had heads four instead of three times as big as those of the smaller men, and jaws nearly five times as big.

The jaws, moreover, of the working ants of the several sizes differed wonderfully in shape, and in the form and number of the teeth.

But the important fact for us is, that though the workers can be grouped into castes of different sizes, yet they graduate insensibly into each other, as does the widely-different structure of their jaws.

I speak confidently on this latter point, as Mr Lubbock made drawings for me with the camera lucida of the jaws which I had dissected from the workers of the several sizes.

With these facts before me, I believe that natural selection, by acting on the fertile parents, could form a species which should regularly produce neuters, either all of large size with one form of jaw, or all of small size with jaws having a widely different structure; or lastly, and this is our climax of difficulty, one set of workers of one size and structure, and simultaneously another set of workers of a different size and structure; a graduated series having been first formed, as in the case of the driver ant, and then the extreme forms, from being the most useful to the community, having been produced in greater and greater numbers through the natural selection of the parents which generated them; until none with an intermediate structure were produced.

Thus, as I believe, the wonderful fact of two distinctly defined castes of sterile workers existing in the same nest, both widely different from each other and from their parents, has originated.

We can see how useful their production may have been to a social community of insects, on the same principle that the division of labour is useful to civilised man.

As ants work by inherited instincts and by inherited tools or weapons, and not by acquired knowledge and manufactured instruments, a perfect division of labour could be effected with them only by the workers being sterile; for had they been fertile, they would have intercrossed, and their instincts and structure would have become blended.

And nature has, as I believe, effected this admirable division of labour in the communities of ants, by the means of natural selection.

But I am bound to confess, that, with all my faith in this principle, I should never have anticipated that natural selection could have been efficient in so high a degree, had not the case of these neuter insects convinced me of the fact.

I have, therefore, discussed this case, at some little but wholly insufficient length, in order to show the power of natural selection, and likewise because this is by far the most serious special difficulty, which my theory has encountered.

The case, also, is very interesting, as it proves that with animals, as with plants, any amount of modification in structure can be effected by the accumulation of numerous, slight, and as we must call them accidental, variations, which are in any manner profitable, without exercise or habit having come into play.

For no amount of exercise, or habit, or volition, in the utterly sterile members of a community could possibly have affected the structure or instincts of the fertile members, which alone leave descendants.

I am surprised that no one has advanced this demonstrative case of neuter insects, against the well-known doctrine of Lamarck.

Jean Baptiste Lamarck
Jean Baptiste Lamarck