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05 - Laws of Variation 05-02 - Use and Disuse of Parts, combined with Natural Selection, Organs of Flight and Vision 20 Kirby has remarked (and I have observed the same fact) that the anterior tarsi, or feet, of many male dung-feeding beetles are often broken off; he examined seventeen specimens in his own collection, and not one had even a relic left.


In the Onites apelles, [?], the tarsi are so habitually lost, that the insect has been described as not having them.
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In some other genera they are present, but in a rudimentary condition.

In the Ateuchus, or sacred beetle of the Egyptians, they are totally deficient.


The evidence that accidental mutilations can be inherited is at present not decisive; but the remarkable cases observed by Brown-Sequard in guinea-pigs, of the inherited effects of operations, should make us cautious in denying this tendency.


Guinea Pig
Guinea Pig

Hence it will perhaps be safest to look at the entire absence of the anterior tarsi in Ateuchus, and their rudimentary condition in some other genera, not as cases of inherited mutilations, but as due to the effects of long-continued disuse; for as many dung-feeding beetles are generally found with their tarsi lost, this must happen early in life; therefore the tarsi cannot be of much importance or be much used by these insects.

05 - Laws of Variation 05-02 - Use and Disuse of Parts, combined with Natural Selection, Organs of Flight and Vision 30 In some cases we might easily put down to disuse modifications of structure which are wholly, or mainly, due to natural selection.

Mr. Wollaston has discovered the remarkable fact that 200 beetles, out of the 550 species (but more are now known) inhabiting beetle, are so far deficient in wings that they cannot fly; and that, of the twenty-nine endemic genera, no less than twenty-three have all their species in this condition!


Several facts, namely, that beetles in many parts of the world are frequently blown to sea and perish; that the beetles in Madeira, as observed by Mr. Wollaston, lie much concealed, until the wind lulls and the sun shines; that the proportion of wingless beetles is larger on the exposed Desertas than in Madeira itself; and especially the extraordinary fact, so strongly insisted on by Mr. Wollaston, that certain large groups of beetles, elsewhere excessively numerous, which absolutely require the use of their wings, are here almost entirely absent;- these several considerations make me believe that the wingless condition of so many Madeira beetles is mainly due to the action of natural selection, combined probably with disuse.


For during many successive generations each individual beetle which flew least, either from its wings having been ever so little less perfectly developed or from indolent habit, will have had the best chance of surviving from not being blown out to sea; and, on the other hand, those beetles which most readily took to flight would oftenest have been blown to sea, and thus destroyed.
05 - Laws of Variation 05-01 - Effects of External Conditions 10 I HAVE hitherto sometimes spoken as if the variations- so common and multiform with organic beings under domestication, and in a lesser degree with those under nature- were due to chance.

This, of course, is a wholly incorrect expression, but it serves to acknowledge plainly our ignorance of the cause of each particular variation.

Some authors believe it to be as much the function of the reproductive system to produce individual differences, or slight deviations of structure, as to make the child like its parents.

But the fact of variations and monstrosities occurring much more frequently under domestication than under nature, and the greater variability of species having wider ranges than of those with restricted ranges, lead to the conclusion that variability is generally related to the conditions of life to which each species has been exposed during several successive generations.

In the first chapter I attempted to show that changed conditions act in two ways, directly on the whole organisation or on certain parts alone, and indirectly through the reproductive system.

In all cases there are two factors, the nature of the organism, which is much the most important of the two, and the nature of the conditions.

The direct action of changed conditions leads to definite or indefinite results. In the latter case the organisation seems to become plastic, and we have much fluctuating variability.

In the former case the nature of the organism is such that it yields readily, when subjected to certain conditions, and all, or nearly all the individuals become modified in the same way.

It is very difficult to decide how far changed conditions, such as of climate, food, &c., have acted in a definite manner.

There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence.

But we may safely conclude that the innumerable complex co-adaptations of structure, which we see throughout nature between various organic beings, cannot be attributed simply to such action.

In the following cases the conditions seem to have produced some slight definite effect: E. Forbes asserts that shells at their southern limit, and when living in shallow water, are more brightly coloured than those of the same species from further north or from a greater depth; but this certainly does not always hold good.

Sea Shell
Sea Shell
05 - Laws of Variation 05-02 - Use and Disuse of Parts, combined with Natural Selection, Organs of Flight and Vision 70 It is difficult to imagine conditions of life more similar than deep limestone caverns under a nearly similar climate; so that, in accordance with the old view of the blind animals having been separately created for the American and European caverns, very close similarity in their organisation and affinities might have been expected.

This is certainly not the case if we look at the two whole faunas; and with respect to the insects alone, Schiodte has remarked, "We are accordingly prevented from considering the entire phenomenon in any other light than something purely local, and the similarity which is exhibited in a few forms between the Mammoth cave (in Kentucky) and the caves in Carniola, otherwise than as a very plain expression of that analogy which subsists generally between the fauna of Europe and of North America."

On my view we must suppose that American animals, having in most cases ordinary powers of vision, slowly migrated by successive generations from the outer world into the deeper and deeper recesses of the Kentucky caves, as did European animals into the caves of Europe.

We have some evidence of this gradation of habit; for, as Schiodte remarks, "We accordingly look upon the subterranean faunas as small ramifications which have penetrated into the earth from the geographically limited faunas of the adjacent tracts, and which, as they extended themselves into darkness, have been accommodated to surrounding circumstances.

Animals not far remote from ordinary forms, prepare the transition from light to darkness.

Next follow those that are constructed for twilight; and, last of all, those destined for total darkness, and whose formation is quite peculiar."

These remarks of Schiodte's it should be understood, apply not to the same, but to distinct species.

By the time that an animal had reached, after numberless generations, the deepest recesses, disuse will on this view have more or less perfectly obliterated its eyes, and natural selection will often have effected other changes, such as an increase in the length of the antennae or palpi, as a compensation for blindness.

Notwithstanding such modifications, we might expect still to see in the cave-animals of America, affinities to the other inhabitants of that continent, and in those of Europe to the inhabitants of the European continent.

And this is the case with some of the American cave-animals, as I hear from Professor Dana; and some, of the European cave insects are very closely allied to those of the surrounding country.

It would be difficult to give any rational explanation of the affinities of the blind cave-animals to the other inhabitants of the two continents on the ordinary view of their independent creation.

That several of the inhabitants of the caves of the Old and New Worlds should be closely related, we might expect from the well-known relationship of most of their other productions.

As a blind species of Bathyscia is found in abundance on shady rocks far from caves, the loss of vision in the cave-species of this one genus has probably had no relation to its dark habitation; for it is natural that an insect already deprived of vision should readily become adapted to dark caverns.

Another blind genus (Anophthaimus) offers this remarkable peculiarity, that the species, as Mr. Murray observes, have not as yet been found anywhere except in caves; yet those which inhabit the several eaves of Europe and America are distinct; but it is possible that the progenitors of these several species, whilst they were furnished with eyes, may formerly have ranged over both continents, and then have become extinct, excepting in their present secluded abodes.


Far from feeling surprise that some of the cave-animals should be very anomalous, as Agassiz has remarked in regard to the blind fish, the Amblyopsis, and as is the case with blind Proteus with reference to the reptiles of Europe, I am only surprised that more wrecks of ancient life have not been preserved, owing to the less severe competition to which the scanty inhabitants of these dark abodes will have been exposed.