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genera having more than the average number of species in any country, the species of these genera have more than the average number of varieties. In large genera the species are apt to be closely, but unequally allied together, 'forming little clusters round certain other species. Species very closely allied to other species apparently have restricted ranges. In all these several respects the species of large genera present a strong analogy with varieties. And we can clearly understand these analogies, if species have once existed as varieties, and have thus originated; whereas, these analogies are utterly inexplicable if each species has been independently created.

{We have, also, seen that it i the most flourishing or ddminant species of the larger nera within each class which on an average vary m tiand varieties, as we shall hereafter see, tend to bgque converted into new and distinct species.)The larger genera thus tend to become larger; and throughout nature the forms of life which are now dominant tend to become still more dominant by leaving many modified and dominant descendants. But by steps hereafter to be explained, theflarger genera also tend to break up into smaller genera? And thus, _t_he__iorm_§ .LfJiiB throughpupthe universe become ’divided into groups subordinate,th smurf“ f “‘ "d d

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CHAPTER III

STRUGGLE FOR EXISTENCE

Its bearing on natural selection—The term used in a wide sense—
Geometrical ratio of increase—Rapid increase of naturalised
animals and plants—Nature of the checks to increase—Com-
petition universal—Effects of climate—Protection from the
number of individuals—Complex relations of all animals and
plants throughout nature—Struggle for life most severe between
individuals and varieties of the same species; often severe
between species of the same genus—The relation of organism to
organism the most important of all relations.

BEFORE entering on the subject of this chapter, I must make a few preliminary remarks to show how the struggle for existence bears on Natural Selection. It has been seen in the last chapter that amongst organic beings in a state of nature there is some individual variability: indeed I am not aware that this has ever been disputed. It is immaterial for u§ whether a multitude of doubtful forms be called species or sub-species or varieties; what rank, for instance, the two or three hundred doubtful forms of British plants are entitled to hold, if the existence of any well-marked varieties be admitted. But the mere existence of individual variability and of some few Well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species""arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life, and of one organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped \ or feathers of a bird; in the structure of the beetle which dives through the water; in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world.

Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most

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cases obviously differ from each other far more than do the varieties of the same species ‘I How do those groups of species, which constitute what are called distinct genera, and which differ from each other more than do ’ the species of the same genus, arise ‘I All these results, as 'we shall more fully see in the next chapter, follow \ from the struggle for life. Owing to this struggle for life, any variation, however slight, and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations ;to other organic, beings and to its physical conditions l of life, will tend to the preservation of that individual, and will generally be inherited by its offspring. The ofispring, also, will thus have a better chance of surviving, for, 01 the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term Natural Selection, in order to mark its relation to man’s power of selection. | We have seen that man by selection can certainly produce lgreat results, and can adapt organic beings to his own uses, hrough the accumulation of slight but useful variations, Elven to him by the hand of Nature, [But Natural Selection,

5 we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man’s feeble efforts, as the works of Nature are to those of Art]

We will now discuss in a little more detail the struggle for existence. In my future work this subject shall be treated, as it well deserves, at much greater length. The elder De Candolle and Lyell have largely and philoso~ phically shown that all organic beings are exposed to severe competition. In regard to plants, no one has treated this subject with more spirit and ability than W. Herbert, Dean of Manchester, evidently the result of his great horticultural knowledge. Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult—at least I have found it so—than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladness, we often see superabundance of food; we do not see, or we forget that the birds which are idly singing round us mostly live on

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insects 01' seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always hear in mind, that though food may be now superabundant, it is not so at all seasons of

e recurring year. Q? should premise that (I usewtfheltellm Struggle for xistence in a large and “m€taphorical‘fsense,._ including dependent-ems? one beingjojnfljanotherJ and including (which is more important) not only the life of the individual, but success in leaving progeny. Two canine animals in a time of dearth, may be truly said to struggle with each other which shall get food and live. But a plant on the edge of a desert is said to struggle for life against the drought, though more properly it should be said to be dependent on the moisture. A plant which annually produces a thousand seeds, of which on an average only one comes to maturity, may be more truly said to struggle with the plants of the same and other kinds which already clothe the ground. The mistletoe is dependent on the apple and a few other trees, but can only in a far-fetched sense he said to struggle with these trees, for if too many of these parasites grow on the same tree, it will languish and die. But several seedling mistletoes, growing close together on the same branch, may more truly be said to struggle with each other. As the mistletoe is disseminated by birds, its existence depends onv birds; and it may metaphoricallyi be said to struggle with other fruit-bearing plants, in order to tempt birds to devour and ‘thus disseminate its seeds rather than those of other plants. In these several senses, which pass into each other, I use for c enience’ sake the general term of struggle for existence. I A struggle for existence inevitably follows rom the high rate at which all organic beings tend to increase. Every being, which during its natural lifetime produces several eggs or seeds, must suffer destruction during some period of its life, and during some season_griccasional year, otherwisefgfilhgjpgififlgmgémmqease; its numbers would quickly become so inordinately great that no country could support the product. Hence, as more individuals are produced than can possibly survive, there must in every case be a struggle for existence; either one individual with another of the same species, or the individuals of distinct species, or with the

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66 STRUGGLE FOR EXISTENCE a

physical conditions of life. {It is the doctrine o? Malthus

applied with manifold force to the whole animal and vegetable kingdoms; for in this case there can be no artificial increase of food, and no prudential restraint from marriage.v Although some species may be now increasing, more or less rapidly, in numbers, all cannot do so, for the world would not hold them.

There is no exception to the rule that every organic being naturally increases at so high a rate, that if not destroyed, the earth would soon be covered by the progeny of a single pair. Even slow-breeding man has doubled in twenty-five years, and at this rate, in a few thousand ‘ years, there would literally not be standing-room for his progeny. Linnaeus has calculated~ that if an annual plant produced only two seeds—and there is no plant nearly so unproductive as this—and their seedlings next year produced two, and so on, then in twenty years there would be a million plants. The elephant is reckoned the slowest breeder of all known animals, and I have taken some pains to estimate its probable minimum rate of natural increase: it will be under the mark to assume that it breeds when thirty years old, and goes on breeding till ninety years old, bringing forth three pair of young in this interval; if this be so, at the end of the fifth century there would be alive fifteen million elephants, descended from the first pair.

But we have better evidence on this subject than mere theoretical calculations, namely, the numerous recorded cases of the astonishingly rapid increase of various animals in a' state of nature, when circumstances have been favourable to them during two or three following seasons. Still more striking is the evidence from our domestic animals of many kinds which have run wild in several parts of the world: if the statements of the, rate of increase of slow-breeding cattle and horses in South America, and latterly in Australia, had not been well authenticated, they would have been incredible. So it is with plants: cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years. Several of

the plants, such as the cardoon and a tall thistle, now most numerous over the wide plains of La Plata, clothing square leagues of surface almost to the exclusion of all other plants, have been introduced from Europe; and there are plants which now range in India, as I hear from

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