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Effect of struggle for existence under unchanged conditions—The effect

under change of conditions-Divergence of character-In insects—In birds—In mammalia- Divergence leads to a maximum of life in each area-Closely allied species inhabit distinct areas-Adaptation to conditions at various periods of life—The continued existence of low forms of life--Extinction of low types among the higher animals— Circumstances favourable to the origin of new species-Probable origin of the dippers—The importance of isolation-On the advance of organi. sation by natural selection--Summary of the first five chapters.

In the preceding chapters we have accumulated a body of facts and arguments which will enable us now to deal with the very core of our subject—the formation of species by means of natural selection. We have seen how tremendous is the struggle for existence always going on in nature owing to the great powers of increase of all organisms; we have ascertained the fact of variability extending to every part and organ, each of which varies simultaneously and for the most part independently; and we have seen that this variability is both large in its amount in proportion to the size of each part, and usually affects a considerable proportion of the individuals in the large and dominant species. And, lastly, we have seen how similar variations, occurring in cultivated plants and domestic animals, are capable of being perpetuated and accumulated by artificial selection, till they have resulted in all the wonderful varieties of our fruits, flowers, and vegetables, our domestic animals and household pets, many of which differ from each other far more in external characters, habits, and instincts than do species in a state of nature. We have now to inquire whether there is any analogous process in nature, by which wild animals and plants can be permanently modified and new races or new species produced

Effect of Struggle for Existence under Unchanged Conditions.

Let us first consider what will be the effect of the struggle for existence upon the animals and plants which we see around us, under conditions which do not perceptibly vary from year to year or from century to century. We have seen that every species is exposed to numerous and varied dangers throughout its entire existence, and that it is only by means of the exact adaptation of its organisation—including its instincts and habits

—to its surroundings that it is enabled to live till it produces offspring which may take its place when it ceases to exist. We have seen also that, of the whole annual increase only a very small fraction survives; and though the survival in individual cases may sometimes be due rather to accident than to any real superiority, yet we cannot doubt that, in the long run, those survive which are best fitted by their perfect organisation to escape the dangers that surround them. This “survival of the fittest” is what Darwin termed “natural selection,” because it leads to the same results in nature as are produced by man's selection among domestic animals and cultivated plants. Its primary effect will, clearly, be to keep each species in the most perfect health and vigour, with every part of its organisation in full harmony with the conditions of its existence. It prevents any possible deterioration in the organic world, and produces that appearance of exuberant life and enjoyment, of health and beauty, that affords us so much pleasure, and which might lead a superficial observer to suppose that peace and quietude reigned throughout nature.

The Effect under changed Conditions. But the very same process which, so long as conditions remain substantially the same, secures the continuance of each species of animal or plant in its full perfection, will usually, under changed conditions, bring about whatever change of structure or habits may be necessitated by them. The changed conditions to which we refer are such as we know have occurred

throughout all geological time and in every part of the world. Land and water have been continually shifting their positions ; some regions are undergoing subsidence with diminution of area, others elevation with extension of area ; dry land has been converted into marshes, while marshes have been drained or have even been elevated into plateaux. Climate too has changed again and again, either through the elevation of mountains in high latitudes leading to the accumulation of snow and ice, or by a change in the direction of winds and ocean currents produced by the subsidence or elevation of lands which connected continents and divided oceans. Again, along with all these changes have come not less important changes in the distribution of species. Vegetation has been greatly modified by changes of climate and of altitude ; while every union of lands before separated has led to extensive migrations of animals into new countries, disturbing the balance that before existed among its forms of life, leading to the extermination of some species and the increase of others.

When such physical changes as these have taken place, it is evident that many species must either become modified or cease to exist. When the vegetation has changed in character the herbivorous animals must become able to live on new and perhaps less nutritious food; while the change from a damp to a dry climate may necessitate migration at certain periods to escape destruction by drought. This will expose the species to new dangers, and require special modifications of structure to meet them. Greater swiftness, increased cunning, nocturnal habits, change of colour, or the power of climbing trees and living for a time on their foliage or fruit, may be the means adopted by different species to bring themselves into harmony with the new conditions; and by the continued survival of those individuals, only, which varied sufficiently in the right direction, the necessary modifications of structure or of function would be brought about, just as surely as man has been able to breed the greyhound to hunt by sight and the foxhound by scent, or has produced from the same wild plant such distinct forms as the cauliflower and the brussels sprouts.

We will now consider the special characteristics of the changes in species that are likely to be effected, and how far they agree with what we observe in nature.

Divergence of Character. In species which have a wide range the struggle for existence will often cause some individuals or groups of individuals to adopt new habits in order to seize upon vacant places in nature where the struggle is less severe. Some, living among extensive marshes, may adopt a more aquatic mode of life; others, living where forests abound, may become more arboreal. In either case we cannot doubt that the changes of structure needed to adapt them to their new habits would soon be brought about, because we know that variations in all the external organs and all their separate parts are very abundant and are also considerable in amount. That such divergence of character has actually occurred we have some direct evidence. Mr. Darwin informs us that in the Catskill Mountains in the United States there are two varieties of wolves, one with a light greyhound-like form which pursues deer, the other more bulky with shorter legs, which more frequently attacks sheep.1 Another good example is that of the insects in the island of Madeira, many of which have either lost their wings or have had them so much reduced as to be useless for flight, while the very same species on the continent of Europe possess fully developed wings. In other cases the wingless Madeira species are distinct from, but closely allied to, winged species of Europe. The explanation of this change is, that Madeira, like many oceanic islands in the temperate zone, is much exposed to sudden gales of wind, and as most of the fertile land is on the coast, insects which flew much would be very liable to be blown out to sea and lost. Year after year, therefore, those individuals which had shorter wings, or which used them least, were preserved; and thus, in time, terrestrial, wingless, or imperfectly winged races or species have been produced. That this is the true explanation of this singular fact is proved by much corroborative evidence. There are some few flowerfrequenting insects in Madeira to whom wings are essential, and in these the wings are somewhat larger than in the same species on the mainland. We thus see that there is no general tendency to the abortion of wings in Madeira, but that it is simply a case of adaptation to new conditions. Those insects

1 Origin of Species, p. 71.

to whom wings were not absolutely essential escaped a serious danger by not using them, and the wings therefore became reduced or were completely lost. But when they were essential they were enlarged and strengthened, so that the insect could battle against the winds and save itself from destruction at sea. Many flying insects, not varying fast enough, would be destroyed before they could establish themselves, and thus we may explain the total absence from Madeira of several whole families of winged insects which must have had many opportunities of reaching the islands. Such are the large groups of the tiger-beetles (Cicindelidæ), the chafers (Melolonthidae), the click-beetles (Elateridæ), and many others.

But the most curious and striking confirmation of this portion of Mr. Darwin's theory is afforded by the case of Kerguelen Island. This island was visited by the Transit of Venus expedition. It is one of the stormiest places on the globe, being subject to almost perpetual gales, while, there being no wood, it is almost entirely without shelter. The Rev. A. E. Eaton, an experienced entomologist, was naturalist to the expedition, and he assiduously collected the few insects that were to be found. All were incapable of flight, and most of them entirely without wings. They included a moth, several flies, and numerous beetles. As these insects could hardly have reached the islands in a wingless state, we must assume that they were originally winged, and lost their power of flight because its possession was injurious to them.

It is no doubt due to the same cause that some butterflies on small and exposed islands have their wings reduced in size. Our common tortoise-shell butterfly is very much smaller in the Isle of Man, and Mr. Wollaston notes that Vanessa callirhoe—a closely allied South European form of our redadmiral butterfly—is permanently smaller in Porto Santo than in the larger and more wooded adjacent island of Madeira.1

A very good example of comparatively recent divergence of character, in accordance with new conditions of life, is afforded by our red grouse. This bird, the Lagopus scoticus of

1 At present I am informed that full-sized tortoise-shells are common in the Isle of Man ; but the late Mr. Edwin Birchall, who had lived there, gave me specimens little more than half the size of ours, which he assured me (about 1865 I think) were the only kind he had then seen in the island.

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