CHAPTER V. LAWS OF VARIATION.

Effects of changed conditions -- Use and disuse, combined with natural selection; organs of flight and of vision -- Acclimatisation -- Correlated variation -- Compensation and economy of growth -- False correlations -- Multiple, rudimentary, and lowly organised structures variable -- Parts developed in an unusual manner are highly variable: specific characters more variable than generic: secondary sexual characters variable -- Species of the same genus vary in an analogous manner -- Reversions to long-lost characters -- Summary.

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 wide 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, etc., 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. Mr. Gould believes that birds of the same species are more brightly coloured under a clear atmosphere, than when living near the coast or on islands; and Wollaston is convinced that residence near the sea affects the colours of insects. Moquin-Tandon gives a list of plants which, when growing near the sea-shore, have their leaves in some degree fleshy, though not elsewhere fleshy. These slightly varying organisms are interesting in as far as they present characters analogous to those possessed by the species which are confined to similar conditions.

When a variation is of the slightest use to any being, we cannot tell how much to attribute to the accumulative action of natural selection, and how much to the definite action of the conditions of life. Thus, it is well known to furriers that animals of the same species have thicker and better fur the further north they live; but who can tell how much of this difference may be due to the warmest-clad individuals having been favoured and preserved during many generations, and how much to the action of the severe climate? For it would appear that climate has some direct action on the hair of our domestic quadrupeds.

Instances could be given of similar varieties being produced from the same species under external conditions of life as different as can well be conceived; and, on the other hand, of dissimilar varieties being produced under apparently the same external conditions. Again, innumerable instances are known to every naturalist, of species keeping true, or not varying at all, although living under the most opposite climates. Such considerations as these incline me to lay less weight on the direct action of the surrounding conditions, than on a tendency to vary, due to causes of which we are quite ignorant.

In one sense the conditions of life may be said, not only to cause variability, either directly or indirectly, but likewise to include natural selection, for the conditions determine whether this or that variety shall survive. But when man is the selecting agent, we clearly see that the two elements of change are distinct; variability is in some manner excited, but it is the will of man which accumulates the variations in certain direction; and it is this latter agency which answers to the survival of the fittest under nature.

EFFECTS OF THE INCREASED USE AND DISUSE OF PARTS, AS CONTROLLED BY NATURAL SELECTION.

>From the facts alluded to in the first chapter, I think there can be no doubt that use in our domestic animals has strengthened and enlarged certain parts, and disuse diminished them; and that such modifications are inherited. Under free nature we have no standard of comparison by which to judge of the effects of long-continued use or disuse, for we know not the parent-forms; but many animals possess structures which can be best explained by the effects of disuse. As Professor Owen has remarked, there is no greater anomaly in nature than a bird that cannot fly; yet there are several in this state. The logger-headed duck of South America can only flap along the surface of the water, and has its wings in nearly the same condition as the domestic Aylesbury duck: it is a remarkable fact that the young birds, according to Mr. Cunningham, can fly, while the adults have lost this power. As the larger ground-feeding birds seldom take flight except to escape danger, it is probable that the nearly wingless condition of several birds, now inhabiting or which lately inhabited several oceanic islands, tenanted by no beasts of prey, has been caused by disuse. The ostrich indeed inhabits continents, and is exposed to danger from which it cannot escape by flight, but it can defend itself, by kicking its enemies, as efficiently as many quadrupeds. We may believe that the progenitor of the ostrich genus had habits like those of the bustard, and that, as the size and weight of its body were increased during successive generations, its legs were used more and its wings less, until they became incapable of flight.

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. 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. 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.

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 Madeira, 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 very 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.

The insects in Madeira which are not ground-feeders, and which, as certain flower-feeding coleoptera and lepidoptera, must habitually use their wings to gain their subsistence, have, as Mr. Wollaston suspects, their wings not at all reduced, but even enlarged. This is quite compatible with the action of natural selection. For when a new insect first arrived on the island, the tendency of natural selection to enlarge or to reduce the wings, would depend on whether a greater number of individuals were saved by successfully battling with the winds, or by giving up the attempt and rarely or never flying. As with mariners shipwrecked near a coast, it would have been better for the good swimmers if they had been able to swim still further, whereas it would have been better for the bad swimmers if they had not been able to swim at all and had stuck to the wreck.

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